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1992-09-01

Species of the Tempskya from Utah

William D. Tidwell

Naomi Hebbert

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BYU ScholarsArchive Citation Tidwell, William D. and Hebbert, Naomi, " of the Cretaceous tree fern Tempskya from Utah" (1992). Faculty Publications. 1445. https://scholarsarchive.byu.edu/facpub/1445

This Peer-Reviewed Article is brought to you for free and open access by BYU ScholarsArchive. It has been accepted for inclusion in Faculty Publications by an authorized administrator of BYU ScholarsArchive. For more information, please contact [email protected], [email protected]. InL 1. Sci. IS3(3):SI3-S28. 1992. e 1992 by The University ofCh icago. All rights reserved . IOS8·S8931921S30 3-OO 27S02.00

SPECIES Of THE CRETACEOUS TREE flRN TEMPSKYA fROM UTAH

WILLIAM D. TIDWELL' AND NAO MI HEBBERT Department o f Bo tany and Range Science. Brigham Young Unive rsity, Provo, Utah 8460 2

Nine species o f the perm ineralized stems of Tempskya were investigated from the upper lower Cre­ taceous Cedar Mounta in and Burro Can yon Pormauons, and the lower Upper Cretaceous Dakota For ­ mation in central and southeastern Utah. Trmpsk}'o joflt'Sii, T. 5tichkar . and T. readii are new and art' differen tiated on the basis of the rad ial orientation of their dorsiventra l stems , their internodal lengths, the lack ofsclerenchyma in the mnerconexof T.j onesii. the three nearl y cont inuous zonesofsclerenchy ma in the inner cortex of T. stichkae, and the com pletely sclerotic inner corte x of T. 'radii. Specimens of T. ionesii and T. minor were collected in growth positiun ncar Cas tle Dale, Utah . which is the first time Tempskya has been collected in this position in North America. Thin were also observed for the li~t time in Tempskya in speci mens of T. ..yomi ngt'tISt'. Tunnels containing three su es o f fecal pellets are common in tissues of stems, . petioles. and epidermal hairs of the false trun ks of Tempskya, ind icati ng that three different grazers fed upon these . Arbuscules of vesicutar-arbuscular (V-A) mycorr hiza l fungi occu r in some roots of the false trunks of T. jo ncui. Specific charac ters of Tempskya and possible environmental circumst ances under which these plants grew are reviewed. and we conclude that they grew under relatively humid conditions in swamps on t100d plains.

Introduction specimens of Tempskya collected from these for­ mations indicate that this fern gre..... abundantly Tempskya. named by Corda (1845), is an un­ in this region at the time the formation s were usual tree fern which lived duri ng the Cretaceous being deposited. period. Specimens of this consist of false trunks composed of a dichotomous branching stem system w hich is surrounded by a thick mat Material and methods of adventitious roots. Although many well-pre­ Specimens of Tempskya are silicified and vary served silicifi ed specimens have been studied, in color from black to gray to red where they had many aspects of these plants remain uncertain. been weathered. Several false trunks have been Becauseof this, much speculation has surrounded collected somewhat intact, although most speci­ Tempskya since it was first reported by Stokes mens are represented by fragments. The majority and Webb (1824) under the name Endogenues of Tempskya specimens from Utah are well pre­ erosa. Since then they have been discovered at served: however, some are not. In these, either manylocalities in the Northern Hemisphere. His­ the stems have disintegrated, leaving cavities in torical accounts abo ut this genus have been pub­ the specimen, or the stem areas have been reo lished by Read and Brown (1937), Ash and Read placed .....ith white crystalline material. The spec­ (1976), and Hlustik (1990). imens are striking aesthetically but are inade­ Geographically, Tempskya is presently known quate for scientific study. only from the North ern Hemisphere, .....here it is Specimens of Tempskya have been collected in restricted generally to the Lower Cretaceous. It growth position near Castle Dale, Utah (locality occurs in England, Ireland, north ern France, Ger­ I), .....hich is rather unusual for these plants (fig. many, Czechoslovakia, USSR, Japan, and ca. 43 2). Specimens from the other Utah localities ap­ localities in the United States. Most of these lo­ pear to have been transported. Many have been calities in the United States are in the Rocky moved only short distances, but they were still Mountain region in such diverse areas as central collected in a horizontal position rather than a Montana, north-central and western Wyoming, vert ical or grow th position. southeastern Idaho, east-central Oregon, south­ Standard thin-sections and acetate peels were western Colorado, northeastern Arizona, south­ made of the and studied with transmitted ern Nevada, northwestern and south.....estern New light. Some of those permineralized with darker Mexico, and Utah (Ash and Read 1976), Our material w ere etched in HF and studied using research on Tempskya has centered on localities reflected light microscopy. in the upper Lower Cretaceous Cedar Mountain Portions of two specimens .....ere chemically and Burro Canyon Formations, and the lo.....er macerated, and the residue of plant material, par­ Upper Cretaceous Dakota Form ation from cen­ ticularly the spores, was isolated for study, tral and southeastern Utah (fig. I). The many Dr. E, M. V. Nambudiri, University of Regina, crushed samples of the carbonaceous (coalified) 'Author for corres ponde nce and reprints. shale associated .....ith the Tempskya to - 20 mesh .\fafluscript receivedOaooer J99 J.. revisedmafluscript received and mounted the samp les as epox.y pellets for January 1991. petrographic analysis.

j)J 5 ' 4 I NTER NATIO NAL J O URNAL OF PLANT SC IEN C ES

Stokes (1952) were tentatively identi fied as T. minor Read and those of Kat ich (195 1) as Tempskya sp. Arnold ( 1958) mentioned observing Tempskya about 9.6 km southeast ofCasric Dale. Utah, near Salt lake City. locality I. The bed in which he saw these speci­ mens lies about 15.2 m below the overlying Da­ \ kota Formation in the Cedar Moun tain Forma­ tion. He tentat ively iden tified them as T. minor. r: We collected fi ve specimens in growth position from th is locality, and curators at the Utah State ; ~e Mu seum of Na tura l History collected another. Castle Dale_a> 6 T heir specimen is a 1.8 m high T. minor. Ferron ~0 Specimens of an unidentified species of Temp­ 170 I skya were reported by McKnight and Rigby (1963) oab from the upper port ion of the Ceda r Mountain (Burro Canyon) Formation northwest of Moab \"'e at locality 6 (fig. I). T idwell and Hebbert (1972. 191 1976) discussed the occurrence of Tempskya in 115 , the Ceda r Mountain Formation from localities Blanding 95 ...... ncar Castle Dale (locality I) and the Moab air­ port, Utah. In a ree valuation of the tempskyas 40 Km mentioned from this formation by Tidwell et aL ( 1976). T. jonesii rem ains a new species in this fig. I (odell map of Utah showing collecting sites for article; specimens called T. wtuteheadii are as­ Tempskya. I, Castle Dale locality; 2, Ferron locality; J, south signed to T. wesselii; and those tentatively re­ oflocalily 2; 4, Yenoweararea: S. Westwater locality: 6, Moab ferred to T. superba are here assigned to T. ..yo­ airport locality; 7. Blue Mesa localit y; 8. Pack Creek locali ty; and 9. Conon wood Creek locality near Blanding. mingense. Tempskya has also been noted from the Burro Canyon and Dakota Formations south­ west of Moab near the La Sal Mountains (locality History of investigations of 8; Read and Brown 1937; Ash and Read 1976). Utah Tempskya Specimens of Tempskya were reported from Lecclltles for Temps1cyo in Utoh the Dakota Formation in San Juan County in Nine species of Tempskya were studied from southeastern Utah by Read and Brown (1937). the Cedar Mountain, Burro Canyon. and Dakota Brown (1950) described a specimen collected by Formations in Utah (fig. I; table I). Th ese consist W.L.Stokesfrom the Dakota FormationinGrand of the three new species. T. jonesii. T. suchkae. County, Utah. as T. knowltonii. However, ac­ and T, readii, under consideration and six spe­ cording to Katich (1952), a reassessment of the cies, T. minor, T. zetierii. T. wesselii. T. wyom­ specimen led Brown to consider placing it in a ingense. T. grandis. and T. superba. described new species, but this was never accomplished . previously either from Utah or elsewhere in the Temp skya was collected during investigations of western United States. the plant s of the Dakota Formation in the locA LITY I. The Castle Dale locality occurs area (Tidwell cr al. 1967; Rushforth and Tidwell 9.6 km southeast of Castle Dale, Utah, near the 1968; Rushfortb 1971). Amateur collectors have con fl uence of Cottonwood Creek and Huntington also collected a large number of Tempskya spec­ Creek which forms the San Rafael River (SWI/4, imens in the Yellowcat area (locality 4) in the SElf4 , S~ . 21, T 19S. R9E). Thi s is ncar Katich's a few miles to the (195 1, 1952) locality and the site where Tschudy west of where Stokes collected his specimen. et al. (1984;USGS KCM D5785) collected the Tempskya has been investigated from the Ce­ palynomorphs for their study. The Cedar Moun­ dar Mountain Formation ncar the type locality tain Formation at this locality is composed of ofthi s formation at Cedar Mountain. Utah (Ka­ bro wn to gray shales and sandstones. It contains rich 1951: Tidwell 1966). Specimens collected by a horizon of nearly coalified material ca. 50 ft - rIgS- 2- 8 Tempskya jofINii sp. nov. Fig.. 2. Three false tru nks of T. jofINii in growth position at the Castle Dale locality. Senior aulhor al kft for scale. fig. 3. Transverse secnon ofthe clesest specim en of T. jo" n ii in fig.. 2 illustrating the stems and roots. Specimen 5045 ( " .3). Fig.. 4. Closeup of stems from fig. 2 showing departing traces . Specimen 5045 ( x 8). Fig.. 5. longitudinal section demonstrating the origin ofthe dkhOiomizing Siems (arro,") from the base of false trunk in fig. 3. Noee .,.....'..0

5

there are atleast three separate branching stems visible in this view, Specimen 5045 (" .75). Fig. 6. Cross sectio n ofthe holotype of T. ionesii. Specimen 5046 (x 1.1). Figs. 7, 8, Cjoseups of sterns of T. ionesii in cross section. Specimen 5046 (7, x 25; 8, x 12). '" INTERNATIONA L J OU RNAL O F PL ANT SC IENCES T.... ' ments, and several mammal teeth (Nelson and TEMPSKYA SPECIES FROM UTAH Crooks 1987). 1..o:AlITY 4. Cedar Mountain Formation in Group I: the Yellowcat area south of Interstate Highway T. kno w/lOni; Seward (Brown 1950; no specim ens of T. know/lOnii were collected or studied by the authors) 70 and west of the Westwater locality (Sec. 9, T. minor Read and Brown T 245, R205). Tempskya jonesii was identified T. zellerii Ash and Read from this area. T. ionesii Tidwell and Hebben sp. nov. LocALITY 5. Westwater locality occurs at the Group 2; same site as the flora reported from the Dakota T. wessettt Arn old NW lf~ , T. wyomingl'nsis Arnold Forma tion by Rushforth (1970, 1971; Sec. T. grandis Read and Brown 17, T19S, R26E). Thirteen specimens of Temp­ T. superba Arnold skya have been collected in this forma tion from T. 51;<:h/uu' T i d ~1I and Hebbert sp. nov. the lower sandstone bed, the lower coal bed , an d Group 3: T. ,rodii Tidwell and Hebben sp. nov. within the ash bed containing the leaf compres­ sions statonidium and Gleichenia. Tempskya jonesii. T. minor. and possibly T. kno .....ltonii (Brown 1950) occur at or near th is locality. locALITY 6. Moab airport locality is situated ( 15.2 m) below the top of the formation from in the low hills ca. 1.6 krn directl y southeast of which several Tempskya false trunks were col­ the Moab airpo rt (Sec. 10, T24S, R20E). Here lected in growth position (Cross et a!' 1975). Py­ tempskyas occur horizontall y and are orien ted in rite occurs in the carbonaceous shales contai ning a northwest-southeast direction in the Cedar the bases ofthese tem pskyas. One specimen also Mountain Formation. although some authors extended through a sandstone lens (Tidwell and (McKnight and Rigby 1963; Ash and Read 1976) Hebbert 1976; Tidwell ct a!' 1976). Specimens of assigned this unit to the Burro Canyo n. McKn ight the angiospcrmous wood Paraphvllamhaxylon and Rigby (1963) noted numerou s fragmented utahense and fronds of the fern Frenelopsis \'ar­ sections of Tempskya at this locality, some up to tens were collected from a nearby site in associ­ 1.5 m long in an argillaceo us siltstone and sandy ation with weathered specimens of Tempskya shale interbedded with pebble conglomerate. The (Thayn et al. 1983). Fronds ofAnemia fremo ntii in situ specime ns were surrounded by a dark gray were also found as compressi ons imbedded in the to black peatlikc material which probabl y rep­ coalified sedime nts around the in situ false trunks. resents the rem ains ofa mat of Tempskya roots. The 500 specimens of Tempskya from this 10· Most of the pieces and the more com plete false cality are represented by T. minor. T. stichkae trunks appeared to have been abraded or partially sp. nov., T. jonesii sp. nov., and T...... esseiii. decomposed prior to burial. This locality con­ LocAuTv 2. The Ferron locality is 15 km east tained 56 specimens assigned to T. stichkae. T. of Ferron, Utah (site 2 in Thayn et a!' 1983; NE If~ , wesseiii (fig. 24), T. wyomingense (figs. 22, 23), SW If~ , Sec. 23, T20S, R8E). At this site, the Cedar T. grandis (fig. 31), and T. ionesii. Mountain Formation consists of coarse, white lcx::A.UTY 7. The Blue Mesa site is sout heast sandstone overlain by channel fills ofyellow con­ of Moab, Utah (NE I/~ , Sec. 16, T28S, R23E). in glomeratic sandstone alternating gray-green shales the upper portion of the Burro Canyon Forma­ an d und erlai n by a dark-green , nod ular weath­ tion. At this locality, abundant specime ns (614) ering shale. Five specimens of T. jonesii occu r of Tempskya occur in a bed of thick, black car­ at thi s locality in association with cycadcoids. the bonaceous shale on top of a cliff composed of conifero us woods Mesembrioxylon stotcesii light tan to brown massive sandstone. Species (Thayn and Tidwell 1984) and Palaeopiceoxylon from the Blue Mesa locality include T. minor, T. thinosus (Tidwell and Thayn 1985 ), and the an­ jonesii, T. readii. T. superba. and T. zellerii. Some giospermous woods Paraphyllanlhoxylon utah­ of the specimens from the Blue Mesa locality, enseand Icacinoxylon piuiense(Thayn ct al. 1983, although bad ly fractured, have been collected 1985). nearly intact from this site. One was over 1.8 m locAuTY 3. This locality is ca. 3.2 km sou th long. A specimen of lcacinoxylon piniense. with­ of locality 2 in SW lf~ , Sec. 26, T20S, R8E. Temp­ ou t growth rings, was associated with these skya occurs above a th in coalified scam ap­ tempskyas. proximate ly 2.5 cm thick in the Cedar Mountain locALITY 8. Thi s is pos sibly the Dakota For­ Formation and over 3.2 m long containing "plat­ ma tion in the La Sal Mountains which Read and anoid" flowers, petrified angiospermous woods, Brown (1937) described as "on the divide south­ two or three types ofornithischian teeth, several west of Winbum 's Ran ch, near the head and on teeth belonging to crocodilians and fish, at least the northwest side of Pack Creek, just west of two different types of dinosauregg shells (Tidwell Mount Peale"(p. 127; possibly Sec. 27, T2 7S, et al. 1983), lizard teeth , tunic carapace frag- R24E) . Read also collected Tempskya ca. 24 km cortex fusing 10 two when connecting with those in the trace. Specime n 5048 ( )( 30). Fig. 2 1, oblique view of the xylem of T. stichkae showing scalariform walls. Specimen 5048 (x 90). Fig. 22, Oblique view of the xylem of T. wyomingense illustrating the scalarifonn walls of the tracheids. Note the absence of parenchyma. Specimen 5049 (x 90). All transverse sections. 518 I N TER N A TION A L JO URNAL OF PL ANT SC IENCES so uth ofthis area (As h and Read 1976). T hey did are solenostelic, dorsiventral, and radially ori ­ not identify the species. ented in the false trunk (figs. 3, 6). The dichot­ locAUTY 9. On Cottonwood Creek ca. 8 km omously bra nched stems are sheathed in adven­ so ut hwest of Blanding, Uta h (S E If~ , Sec. 24, T37S, titious roots which generally parallel the stems R21 E). Although the sha le in which th e tem p­ (fig. 5) and, in so me specimens, in vade th e stems skyas occur is assigned to th e Burro Canyon For­ in the older portions of the false trunks. marion. the strata more closely resemble th e Cedar The epide rm is is compo sed of ta ngentially Mo untai n Formation. The on ly species observed elongated cells whic h have masses of very long by the au thors from th is locality is T. minor. epiderm al hairs extending from th em. A cuticle, 3 urn thick, covers the epidermis. Systematics (figs. 2- 40, table 2) The cortex consists of three parts (figs. 7, 8). FAMILY. Tempskyaceae T he o uter cortex consists of la rge, tangentially GENUS. Tempskya Co rda elongated parenchyma cells. T ran sversely, this SPECI ES. Tempskya Jonesii sp. nov. (figs. 2-12) tissue is 3- 4 cells wide . Homogeneous sc1erenchyma to us tissue forms the m idd le cortex, and the inner cortex is com ­ D IAG NOS IS. False trun k. upward of 30 em in posed of only parenchyma cells (figs. 9, 10). Th e dia meter. radially sym me trica l; individual stems latter tissue is 12- 14 cells wide. Its cells are gen­ dorsiventral. z-ranked, 4-4.5 mm in diameter; erally filled with dark conten ts. Because of its dichotomizing; solenostelic; inte rnodes very long, parenchymatous nature, the inner cortex is often II mm, generally showing o ne, rarely two, leaf not or on ly partiall y preserved in many speci­ traces in cross section; ou tercorte x 3-4 cells wide mens. transversely, contai ns ta ngentiall y elongated pa­ The endoderm is, altho ugh poorly preserved, renchyma cells; m iddle cortex 12-1 5 cells acro ss, appea rs as a single layer of dark cells o n each side hom ogeneou s, sclcrcnchymatous, 12-1 5 cells of the xylem stran d and en sheathes the pcricycle. across; inner co rte x 6-12 cells in wid th, paren­ phloem, and xylem shea th. The pericyc1 e appears chy matous, filled with dark co ntents; xylem with­ to be upwa rd of two cells th ick and th e phloem ou t parenchyma: pith consisting of two zones, vari es from on e to three cells in width. outer zone parenchymatous, inner lone sclcren­ The xylem is 10 (rarely 13) cells thick and con­ chy matous. sists only of angular, sometimes flatte ned and di storted, scalariform tracheids (fig. 12). The pro­ REPOS ITORY. BYU 5046 (holotype). 5045, toxylem is exarch and com posed of small tra­ 505 1 (paratypes). cheids with spi ral thicken ings. A two-zoned pith with parenchyma cells next to the inner endo­ locALITY. 1- 7. dermis and sclerenchyma constituting the m iddle AGE. late Early Cretaceo us (Albian) and ear­ portion is enclosed with in the vascular strand of ly Late Cretaceo us (Cenomanian). the stem. Leaf traces arise in two ranks from the stem. HORIZON. Cedar Mountai n. Burro Canyon. T heir place of origin is marked by a th inning of and Dakota Formati ons. the stele. The xylem is 3-6 tracheids wide at this po int. In transverse section, the lea f trace begins ETYMOLOGY. T he specific name honors Stan­ as an outwa rd bu lging of the vascular tissues of ley Jones of Castle Dale, Utah, for showing us the stem which for ms the beginning ofa gap. The th e Castle Dale locality. parenchy ma and sclerenchyma of the pith also bulge o utward and fiJI the cen ter of the gap. As the gap expands outward, it expands tangentially Description as well, becoming somewhat pea r shaped. The False trunks ofthis species vary fro m round to lea f trace separates from the ste le first on one side flattened (fig. 2). The latter sha pe, however, re­ and at a higher level on the othe r, The three zones sults largely from uneven weathering. The ho­ of cortical tissue encircle the lea f trace as it passes lotypc contains upward of 270 stems in a cro ss outwa rd; and parenchyma and sclcrenchyma of section nearthe base ofthe false tru nk . The ste ms th e pi th occupy the adaxial curvature of the trace.

fIgS. 16-22 Fig. 16. False trunk or Tempskya snchkae sp. nov. in transverse section. Specimen 5048 (x 1). Fig. 17. Stem with departing lear tract' in raise trunk or fi g. 16 ( " 10). Fig. 18, Enlargement or outer pith or a stem or T. stichkae showing the three bands or sclercnchyma (arrows) with the two outermost fusing together in certain areas. Specimen 5048. I ( " 30). Fig. 19. Enlargement or the inner cortex or T. sncnkae illustrating the three bands or sclerenchyma (arro ws) in this tissue. Specimen 5048. I ()( 20). Fig. 20. Lear ' race or T. .sucnkao departing from stem. Note the three sclcrenchyma bands or inner figs. 9-1S Figs. 9, 10, 11, Parenchymatous inner cones (errowsj in transverse sections or a ponion or a stem or Tempskya jonNii. Specimen 5046 (9. " 25; 10. " 75; II , " 90). Fig. 12. Longitudinal view or metaxylem elements or the xylem or a stem or T. jonesu. Specimen 5046 [ x 90). Fig. 13, Cross section or a stem or T. readii. Specimen 5047 (" 8). Fig. 14, Closeup ora stern or T. readii in transverse section showing the sderenchymatous inner corte" (arrows). Specimen 5047 ( " IS). Fig. 1S. Enlargement ofthe inner cortex or a stem or T. readi; illustrating the sclerenchyma (arrows) in the inner cortex. Specimen 5047 (" 90). >20 I N T ER NATION AL JOURNA L OF PLANT SC I ENCES

T he Ccshaped trace enters the base upon and filled with dark contents. The middle co rtex leaving the stem . The xylem strand in both the consists of homogeneous, sclcrenchymatous tis­ leaf traces and pctiolar vascular strands is thin in sue of cells with small lumens filled with da rk the middle(I-3 tracheids in width) and thickened contents and thick walls which are often not pre­ laterally. The petioles are sho rt and often are in­ served. Parench ymatous tissue containing three vaded by roo ts. Ad ventitious roots arise from the thin ( 1-3 cells wide), more or less continuous stele opposite the leaf traces. zones of sclere nchyma consti tutes the inner cor­ tex (fig. 19).Genera lly, the sclero tic zo nes are Comparison separate from one another, but occasionally T he lack ofparenchyma in the xylem and scle­ strands of sclerenchyma connect the innermost renchyma in the outer pith and inner cortex in and middle zone together. T he inner zone is 7­ Tempskya jonesii is similar in T. minor and T. 9 cells wide, and the outer zone is 8-10 cells thick. knowltonii. However. longer internodes and a ra­ Although generally poorly preserved, the Cas­ dial arrangement of the Siems in the false trunk parian strips are seen on some cells ofthe single­ of T. jonesii distinguish it from these species. layered endodermis. and a single-layered peri­ cycle has cells 25.um in diameter. Phloem, where SPEO ES. Tempskya stichkae sp. nov. preserved, appears as patches 1-3 cells wide. The (figs. 16-2 1) xylem shea th is not preserved. Angular, som etimes distort ed, scalariform tra ­ DIAGNOSIS. False trunk. over 17 em in di­ chcids with out parenchyma compose the xylem amete r, radi ally symmetrical; stems dorsiven­ strand (fig. 21). No definite protoxylem could be tral, solenostelic, 6-8 mm in diameter; leaves iden tifi ed. 2-ranked; internodes intermediate, 5-7 mm long The pith consists ofa sclerotic center and pa­ with 1-2, rarely 3, leaf traces present in cross renchymatous outer layers , with three zones of section; epidermal cells filled with dark contents sclcrenchyma similar to those in the inner cortex with epidermal hairs attached; outer cortex thin, occurring next to the inner endodermis (fi g. 18). 4- 5 thick-walled cells wide; middle cortex ho­ leaf traces arise in two ranks from the stem mogeneous. sclerenchymatous. 20-25 cells wide; and develop essentially in the same manner as inner cortex parenchymatous with 3 thin, dis­ those in Tempskya jonesii. Th e three zone s ofthe con tinuous zones of sclerenchyma more or less inner cortex continue around the outside of the evenly spaced an d occasionally connected; xylem leaf trace, but where they connect with the three parenchyma absent; pith, ha ving a sclerotic cen­ zones of sclerenchyma from the pith , the middle ter, parenchymatous near inner endoderm is, con­ and outer zones fuse, resulting in only two zones taining three zones of sclerenchyma similar to of this tissue lining the adax ial concavity of the that of inner cortex. leaf trace and pctiolar vascular strand (fig. 20). Petiol es in this species extend 20 mm from their REPOSITORY. BYU 5048 (holo type). sterns before the y are destroyed by roots. Roots originate from the stele opposite the leaf traces. l.ocAUTY. I, 6. Com parison AGE. Late Early Cretaceous (Albia n). Speci mens of Tempskya stichkae ap pear to be similar to T. supcrba in general intern al structure, HORIZON. Cedar Mountain and Burro Can­ alt hough they differ in stem size. The stems are yon Formations. 6- 8 mm in diam eter, but the departin g leaf traces cause them to a ppear much larger. Th e outer pith ETY MOLOGY. T he specifi c epi thet honors and inner cortical layers in the holotype of this James Stichka of Walnut Creek, California, for new species are well preserved and contain three his continued scientific interest in Tempskya. irregular strands of sclerenchyma. rather than one irregular band as in T. superba (Arnold 1958), or Description two as in T. wyomingense(Arnold 1945). In gen­ Known false trunks of this species are incom­ eral. it is unlike any other species of Tempskya. plete (fig. 16). Th e holotype is 18 cm across with radially oriented dichotomously bran ching stems. SPECI ES. Tempskya readii sp. nov. A poorly preserved epidermis with numerous (figs. 13-1 5) attached epiderma l hai rs and three zones of cor­ DIAGNOSIS. False trunk, radially symmetri­ tical tissue form the outer portion of the stem . cal; stems dorsiven tra l, sole nostelic, 5- 7 mm in Cells of the epidermal hairs are 50 «m wide and diameter; leaf traces 2-ranked; internodes inter­ 150 «m long near their bases. The hairs are very mediate in lengt h, 1-2lcaftraces, occasionally 3, long and also occur on the petioles and roots. present in cross section: outer cortex 6-8 cells Cells of the thi n outer cortex are 50-60 «m across wide; middle cortex homogeneous, sclerenc hy- TIDWELL & H EBBERT_ TEMPSK Y A F RO M U TA H 521 matous, 11- 13 cells wide; inner cortex complete­ tened apex. The annuli are extend ed outward from ly sclerenchyma tous; xylem parenchyma lacking; a short stalk to which they are connected. The pith sclerotic both middle and near inner endo­ stalks are 50- 60 s m wide x 70 urn long, and the derm is. a nnuli are 655 sm long by 16 sm wide. They consist ofcclls 4- 6.urn across with cell walls vary­ REPOSITORY. nY U 5047 (ho lotype). ing from I to 4 «m in thickness. T he annuli ap­ pear to be of the vertical type. No spores are LocALITY. 7. prescnt. A detached annulus present among the roo ts AGE. Late Early Cretaceous. in a false trunk of T. wessetii (fig. 38) is very similar to those reported in this species from HORIZON. Burro Canyon Formation. Waya n, Idah o (Andrews and Kern 194 7).

ETYMOLOGY . The specific epithet honors Animol o

..':; .. . o · •29 fIgS. 23-30 Fig. 23, T ransverse section of a stem of Tempskya wyomingense. Note the sclerenchyma (arrows) in its inner cortex. Specimen 5049 ( x 8). f ig. 24, Cross section of a stem of T. wessetii with departing leaf traces. Specimen 5050 (x 15). Figs. 25-27, Transverse sections of leaves in T. wyomingense. The palisade layer is prominen t, whereas the mesophyll has been crushed (arrow). Specimen 5049 (25. x 30; 26, x 60; 27, x 240). Fig. 28, Section through a sorus that is sessile on a leaf with the annuli (arrows) of seve ral sporangia still attached. Specimen 5049 ()( 240). Fig. 29, A portion of cuticle that remained a fier macerating a stem ofT.jcmn;;. Specimen 50S I (x ISO). Fig. 30, Framboidal pyri te. macerated from carbonaceous shale ( x 4(0). m . ,

,..

••• • : ~

39

Flgs. 31-40 Fig. 31, Transverse section of Temp skya grandis. Specimen 5052 (x 1.5). Fig. 32, Cross section of T. zellem. Specimen 5053 (x 2). Figs. 33, 34, Fecal pellets in tunnels in the epidermal hairs of two stems of T. wyomingcnse. Specimen 5049 (x 25). Fig. 35, Arbuscules of V-A mycorrhiza l fungi in cells in a root ofT. wyomi ngense. Specimen 5049 (x 250). Figs. 36. 37. Unidentified germinating fungal spores from maceration of false tru nk of T. jonesii. Specimen 5051 ( x 4(0). Fig. 38, Ann ulus of a sporangium in T. wesselii. Specimen 5050 ( x 25). Figs. 39, 40, Spores from the maceration of a ponion of a false tru nk of T. joneui. Specimen 5051 (" 350).

m 5" INTERNA TIONA L JOU R NAL OF P LANT SCI ENCES ctrati ng the harder tissues as evidenced by the (193 7), Arnold (1945 , 1958), Chandler (1968), general absence of fecal pellets in the pith of the Ash and Read (19 76), and Nishida (1986). Per­ stems. unless the stems were first split apart by haps the basic criterion is whe ther a species has invading roots. Designation ofany particular an­ radial or dorsiventral symmetry (Seward 1924). imal to these pellets is pro blematical beca use di f­ Although the idea of dorsivem rality has been ferent ani mals in diverse groups produce the same viewed with skepticism by some (Andrews and or similar pellets (Hentzschcl ct al. 1968). Kern 194 7), others have deem ed it to be valid (Read and Brown 193 7; Ash and Read 1976; Fungol activity Nishida 1986). Tempskya jonesii is very similar Hyphae resembling the dichotomous bran ch­ to T. knowltonii. and the major difference be­ ing or arbuscules of vesic ular-arbuscular (V-A) tween the two is that the symmetry for T. jonesii mycorrhizal fungi occ ur in the cortices of some is radial, whereas that of T. know/tonii is dorsi­ roots in these false stems, particula rly Tempskya ventral. The specimens of T. minor included in jonesii (fig. 35). The branches appear to become this study appear more radial than dorsiventral; progressively smaller in size with each level of otherwise they are anatomically simi lar to the branching. The genus to which they can be as­ type. Dorsiveru ral trunks were postulated to have signed is problematical. but living forms are grow n alo ng the ground and then eventua lly to known to occ ur in thalli , rhizom es, or roots of have grown upri ght (Seward 1924). Read (1939) (Bonefantc-Fasolo 1984). Several forms of thought perhaps they grew as Hanas or vines lean­ living V·A mycorrhizas may also infect the same ing again st rock outcrops ofsteep cliffs or climb­ plant. All living members are placed in the family ing on arborescent plants. We did not see any Endogo naceae ofthe orde r Mycorales in the class indication of this occ urring, and in our opinion, Zygomycetes (Hall 1984). In living forms, these it never did. Among tree ferns, there are species mycorrnizas represent an extreme form of ad­ illustrating these different growth patterns. Ci­ aptation to a symbiotic mode oflife. It is uncer­ bonum barometz. for example. has a creeping tai n what contribution these fossil forms may have habit of its stem which differs from the upright made to the Tempskya roots. They may have species ofthe genus (Ogura 1927). However, un­ stimulated growt h. helped in nutrient uptake, im­ like T. knowltonii. the stele ofC barometz retains proved wa ter tra nsport, and possibly increased its radial sym metry. the resista nce to pathogens as they do in modern Th e arrangemen t of the tissues in the pith and roo ts (Jackson and Mason 1984). cortices of the stems, particularl y in the inn er Read (1983) concludes that living V-A mycor­ cortex . is considered to hav e systema tic value rhizas do best in areas of high mineral nitrogen (Read and Brown 1937; Arnold 194 5, 1958). Ash availability with low pol yphenol content in low and Read (19 76) subdi vided the western North latitudes and at low alti tudes. Some species, how­ American species into two broad gro ups based ever, do occur at high altitudes and in the most upon the character of this tissue: one group was acid moor and bog soils. They develop the grea t­ cha racterized by a simple three-layered type of est in moist temperate and tropical lowlands where co rtex with an inner parenchymatous cortex. and plants con taining V·A mycorrhizal are dominant the othe r included species with mixed tissues of in the vegetation (Ha rley and Smith 1983). T his parenchyma and sclerenchy ma in their inn er cor­ may have been the case in their association with tices. Like Ash and Read (1976). we consider this Tempskya as well. to be a valid character, and T. jonesii is hereby Nu merous fungal spores are also present in the placed in the fi rst grou p and T. stichkae in the maceration residue of a false trunk of T. jonesii second. However, the inner cortex and the out er (figs. 36, 37). It is uncertai n whether any of these zone of the pith of 1'. readii are completely scle­ spores were prod uced by the V-A mycorrhizal rotic in its stems. The completely sclerotic inner fungi. No hyphae are associated with the spores. cortex of T. readii does not allow this species to be assigned to the other two groups. T herefore , a third gro up con taining only sclerenchyma in its Discussion inner cortices is proposed for T. readii. Twelve species of Tempskya ha ve been de­ The center ofthe pith is sclerotic in all species, scribed from No rth America, eight, possibly 10, except T. reesidei, which has a parenchyma tous from Europe , most of which are im perfectly center, and 1'. readii. wh ich ha s a co mpletely scle­ known, and one from Japan. The latter species rotic pith. In gene ral, the ou ter zone of the pith (1'. iwatensis) was repo rted from U pper Creta­ is similar to the inner cortex as to the disposition ceo us strata (Nishida 1986) rat her than being of of sclercnchyma and parenchyma in that partie­ Early to mid-Cretaceous age as were the others ular species ofTempskya. Tempskya wessetii and (Ash and Read 1976). T. wyo mingrnse, however. differ in lacking scle­ Characters delimiting the species in Tempskya renchyma in the outer layer of their piths at the (table 2) have been reviewed by Read and Brown sam e time they have sclercnchyma in the ir inner TIDWELL & H EB BER T - T EM I'SK YAF ROM UTAH cortices. Th is difference is also a valid specific ruptured. and it is possible to observe only the character. outer portion of the ruptured stem, portions of Stem diameter has been used to a limited extent the pith. leaf traces. and other internal structures for specific separation (Arnold 1958), but Chan­ of the stem. Invaded stems which have not rup­ dler (1968) considered stem size alone not to be tured are expanded until they are much larger sufficient reason for separation, and. in general, than normal, and the major portion ofthe inter­ this may be true. She stated that stems varied nal structure has been replaced by roots. The ar­ considerably in size in her specimen depending rangement ofthe stems within the false trunks in upon their position within the false trunk. We the Utah specimens of T. minor also appeared to have studied many nearly intact specimens and be more radial than do rsiventral. have found very little variation in stem size from A variety of'fungal (fi gs. 36, 37) and fern spore the bases of the false trunks to near their apices. types have been observed in the macerated ma­ In other words, if they are large. such as those in terial from the false trunks of T. jonesii. Most of T. superba. they remain large, and if they are the fern spores (figs. 39, 40) are similar to those small. such as those in T sjonesii, they stay small. illustrated by Andrews and Kern (1947, text fig. The correlation between stem diameter and in­ 8), and although not conclusive, the number of ternodal length used as a specific character by these spores present suggests they are most likely Read and Brown (1937) has been largely dis­ the spores produced by Tempskya. These spores. counted (Ash and Read 1976). The internodal however. diffe r from those described by Boodle length itself is valid as a specific character. The ( 1895) from a specimen ofT. schimperii collected relative length ofthe internodes in Tempsk va has on the Isle of Wight. The spores from the Utah been derived by using the number of visible leaf specimens are smoot h walled rather than being traces in a cross section. However, if the length conspicuously sculpt ured with long bars as are of the internodes were determined by actual mea­ those from T. schimperii. surement ofl ongitud inal sections ofthe stem, the Temp skyas collected in growth position at the results would likely be more accurate. Castle Dale localit y (locality 1) were separated Using the amount of parenchyma in the xylem from each oth er by ca. 1.2 m and were either to distinguish species of Tempskya has been ques­ truncated or tapered to a fl att ened top. None had tioned (Arnold 1958: Ash and Read 1976). Among preserved apices. These specimens were embed­ conifers, the presence of paren chyma in the xy­ ded in undistu rbed carbonaceo us shale contain­ lem are actually the tips of overlapping tracheids. ing pyritic sulphur, which im plies that they in­ The systemic value of the persistence of peti­ habited a reducing, swampy environment similar oles in the false trunks away from the stems is to some of those in the Mississippi River Delta Questionable. Petioles are destroyed by root s and. or the Florida Everglades. The abundant sulphur therefore, their number would vary as to their minerals in these carbonaceous shales occur position in the false trun k. They are generally mostly in the form of framboi dal pyrite which is invaded by root s in older porti ons of the trunk formed in sulturitic environments during early and appear to have ceased functioni ng at the time diagenesis. Although mari ne peats have higher of root invasion. sulphur content than freshwater peats (Casa­ Roots are essentially the same in all species of grande 1987), pyrite is most abundant in peats Tempskya and therefore of no definite taxonomi c formed under brackish water environments (Co­ value. In some species, such as T. stichkae. there hen et al. 1987). Other abundant minerals in these arc small differences. The proportion of larger sediments include limon ite. clay, and carbonate. roots to smaller ones is lower in this species than Vitrin ite macerals in these shales are generally in the ot hers where root s are closer to being the pitted and occur along with fusinites. Pyrofusin­ same size. The roots of T. zellerii also have a ites. with higher reflectance, are frequently en­ tendency to become distorted and angular, which countered in the coalified materia l in these car­ does not occur in the other species. Root s of all bonaceous shales and probably resulted from species, except T. zellerii. encountered in this charri ng of peat swamp vegetation d uring occa­ study have an abundance of attached epidermal sional surface fi res (Stach et al. 1982, pp. 272­ hairs as noted by Nishida (1986). These hairs are 274). It is unclear whether such fires also affec ted essentially the same as those produced by the a forested area near the site of deposition ofthese epidermi s of the stem and, in the case ofT. such­ tree fern s (Namb udiri, personal communication kae. the petioles. Epidermal hairs are not present 1987). on the petioles in the other species. The epidermal Based upon the abundan ce of Tempskya spec­ hairs in T. stichkae are wider at their bases than imens and their wide geographic distribution in those of other species. Utah and adjacent states, these swamps must have The false trun ks of T. minor do not contain been fairlycommon du ring the time ofdeposition many stems, and most of those observed have of the Cedar Moun tain and Burro Canyon For­ been invaded by roots. These invaded stems have mations and other forma tions of equivalent age. ,,, I NTERNA TIONAL JO URNAL OF PL ANT SCIENCES

Tobio 2 Table 2 (Continued)

a ) False trun k dorsrvenua t Xylem txan;h. Stems approximately 25-3.5 mm in dia m- I. Individual stems of false: tru nk medium to large (5- 15 eter. Tempskya kflOw/to"ii mm) in diameter. Inner parenchymatous layer of the b) False trunk rad ially sym metrical. Xylem ex­ co rtex co ntaining connnucus and/or discon tinuous arch or possibly slightly immersed in some bands o f sclerencnvma. False: trunks ha ve radial sym­ specim ens. Xylem ring containing much pa­ met ry. renchyma (?). Stems 6-7 mm in diameter. A. Inner parenchymatous layer of cortex and exterior Tempskyo rouica of pith contain a single discontinuous and irregular band ofsclerenchyma. Internodes shon 10 med ium Note. In earlier reports on Tem pskya (Read and Brown permitting ove rlapping (2- 5) of leaf bases. 1937; Ash and Rcad 1976) efforts were mad e to summarize I. Indi vid ual stem s medium, 4- 6 rom in dia meter. the distinctions of the better-known species of Tempskya. Tempskya grandis Accordingly. a modified key has been prepared which includes 2. Indi vid ual sterns large, 10- 15 mm in diam eter. the better-known and new species of the genus proposed in Tempskya superba th is pape r (modified from Ash and Read [1976) with per­ B. Inner parenchymatous layer ofcortex con tains two mission crs . R. Ash). bands of sclerenchvma. separated by a continuo us or d iscontinuous band of parenchyma . Outer band of scjerenchyma is continuous. and the inner band Tempskya associated with the coals in the Dakota may be either con tinuous or discontinuous. Exterior Formation reinforces the concept that they grew of pith does not contai n a band of sclerenchyma, Internodes medium to long, permitting very ~ i gh t under swam py conditions. Along with Anemia 10 slight overlapping ( 1-3) of leaf bases . fre montii. specimens of Tem pskya occur in the I. The two bands ofscjerenc hyma in the inner pa­ Ceda r Mounta in Formation with fossil woods of renchymatous layer of cortex conn ected locall y Paraphyilanthoxylon tuohense and Icacinoxykm by strands of sclerenchyma, giving an impres­ piuiense, which lack growth rings, and Mesem­ sion o r-tstancs" o f parenchyma surrounded by sclerenchyma . lmem odes med ium in length , brioxylon stokesii with indistinct rings. Lack of perm itting slight o verla pping (2-3)o flea f bases. rings is considered to be evidence for equable Pith pare nchymatous. Tempskyu reesidei year-round temperat ures and rainfall (Crebcr and 2. The two bands ofsclerenchyma in the inner pa­ Cha loner 1985;Crabtree 1987) in humid areas renchymatous layer of cortex completely sepa­ rated from each other by a cont inuou s band of (Crebcr 1977). Paraphyllanthoxylon idahoense parenchyma. Ou ter pith lacks scleren chyma. Spackman (194 8) is associated with T. wesselii a) Internodes long, permitting very slight over­ in the Lower Cretaceous Wayan Formation in lapping (1- 2) ofleaf bases. Ind ivid ual stem s southeastern Idaho. This angiospermous species 4-5 mm in d iameter. Tempskya wt."SSeIii also lacks growth rings. The nearest living rela­ b) Internodes medium in length, permitting slight overlapping (2-3) of leaf bases. Indi­ tives ofthese angiospermous woods are found in vidual stems 6-8 mm in diameter . subtropical or tropical regions (Thayn et al. 1985). Tempskya ",yomingeruis Conifer wood having ind istinct growth rings as­ 3. Inner parenchymatous layer of cortex and ell­ signed to Cupressinoxylon sp. (Andrews and Kern terior of pith contains three bands o f scjeren­ chyma separated by cont inuous or di scontinu­ 1947) and leaves ofAnemia fremonni and Gtei­ ous band s of parenchyma. Outer band contincous chenites coloradensis have also been collected as­ and inner bands either continuous or d iscontin­ sociated with Tempskya from the Wayan For­ uous. Internodes medium (2-3). mat ion. These leaf forms have been related to the Tem pskya stichkae modem genera A nemia and Gleichenia which are 4. Inner layer of cortex and pith completely scle- rot ic. Tem pskya readii tropical to warm subtropical in their present dis­ II. Indi vidual stems offalse trunk small to med ium (2.5­ tribution (Andrews and Kern 1947). 8.0 mm) in dia meter. Inner parenchymatous layer of Tem pskyas in the Dakota Formation grew with the con ex does not contain contin uous or discontinuous ma ny differen t forms of angiosperms and ferns laycn of sclerenchyma. False tru nks ha ve either dor­ sivent ral or radial sym metry. (Rushforth 1971), particularly Gteichenia and A. Ind ividual stem s and steles angular in transverse .\latonidium. whose nearest living relative Ma­ section, approximately 3-8 mm in diameter. Inter­ tonia grows on the Malay Peninsula and Borneo nodes relatively long, perm itting only ~ igh t ove r- (Seward 1899). lapping (1- 2) ofleaf bases . Tempskya zellcrii The presence of growth increments in speci­ B. Individual stems and stdes ro und to subro und in transverse section . Internodes long, perm ittin g slight mens of Paleopiceoxylon thinosus, which have overla pping (2- 3) or leaf bases. been collected in association with Tempskya and I. False trunks doniventral. somet imes approach­ the fossil woods without rings at the Ferron lo­ ing radial. Stems approximately 2.~ 3 .5 mm in cality, can be explained in three ways. Thi s plant diameter. Tempskya minor 2. False trunks rad ial. Stems approximately 4-4.5 may have lived on an eda phic site with poor water mm in d iameter. Tempskya jonesii supply in a tropical or subtropical region as in­ 3. False tru nks doniventral or radiall y symmetri­ dicated by the other plants or it may have lived cal. Stems approximately 25- 7.0 mm in di­ in a wetland forest. marsh, or bog at a low ele­ ameter. vation with long periods of inundation phase which will also produce growth rings (Worbes T IDWEL L & H EBBER T - TEM PSKYA FROM UTA H m

1989). A third possibility is that P. thinosus may bility of more than one plant being involved in have been transported from higher elevatio ns the formation of each false trunk. Gra nting this wherecyclic climatic conditio ns very likely could possibility. then the potential exists for more tha n have occurred and reposited in a lowland area . one species of stems being present in the same A habitat for Tempskya suggested by Andrews false tru nk. However, there is no indication that and Kern (194 7) and Nishida (1986) is a humid this ever happened. It is more likely that the ga­ low to high montane forest in subtropi cal or even metophyte of Tempskya was broad and vascu­ warm temperate climate. Modem tree ferns re­ larized. and that the sporeling stage gave rise to Quire high humidity for continuo us growth such the initial stems in the false trunks. The game­ as in tropical to subtropical South America. where tophyte. being composed of thin -walled cells. cloud forests rich in fern s are well developed on would have eventually disintegrated and added Andean slopes (N ishida 1986)_They also grow nutrients to the trunks. Since the roots of Temp­ at lower eleva tions in climates characteristic of skya appear to have invaded almost every­ New Zealand, Australasia, and Malaysia. How­ thing else, they very likely destroyed their own ever, tempskyas from Utah appear not to have gametop hytcs as well. grown in montane areas but, rather, in swamps on fl ood plains. Since evaporites are not associ­ Acknowledgments ated with Tempskya localit ies a lower cvapora­ For contributing specimens and locality data. tion rate resulting from high humidity (Parrish et we wish to thank Mr. and Mrs. Harry Cleaveland, al. 1982; Nishida 1986) was likely required by Stan Jones, and Mr. and Mrs. Frank Lemon, Dr. Tempskva. This concept strongly contrasts with D. A. Medlyn, Paul Rechten, Dr. J. Roth, A. D. the proposition that much ofthe Cedar Mountain Simper, Dr. W. L. Stokes. J. B. Sanchez, Dr. W. Formation was semiarid. with caliche formi ng Shad ish. James Stichka, Leslie Smith, Dr. G. at the time it was being deposited (Nelson and Tbayn, and Mrs. Marian Whitehead. Special Crooks 1987). than ks are extended to Dr. Cha rles Beck of the Seward (1924) considered that Tempskya grew University of Michigan and Dr. Francis Hueber either partially or completely beneath the soil in of the Smithsonian Institution for lend ing holo­ its beginning stages, later becoming erect or ver­ types of the various North American species of tical. There was no indication that this happened Tempskya. to Dr. E. M. V. Nambudiri of the in the false trunks found in growth position at Energy Research Unit ofthe University ofRegina the Castle Dale locality (locality I). The sedi­ for analyzing the carbonaceous shales and coal­ ments aro und these trunks were compact, and if ified material at the Castle Dale site, to Dr. S. R. any horizon tal false trunks had occurred below Ash of Weber State University and to Dr. A. T. the vertical ones, the n there should have been Cross of Michigan State University for reviewing some indicat ion of their occurrence. Bases ofthese the manu script, and to Dr. Lee Parker of Cali­ vertical false tru nks had from three to nine lobes, forn ia Polytechnic State University for his assis­ with each lobe contain ing a stem that divided tance with some of the research on this fossil dichotomously upward. Th is implies the possi- material.

titercrure cited

Andrews, H. N., and E.M. Kern. 1947. The Idaho tem p­ Chandler. M. E. J. 1968. A new Tempskva from Kent. Br. skyas and associated fossil plants. Ann . Mo. Bot. Gard. Mus. (Nal. Hist.) uuu.. Grot 15(4):169- 179. 34(2):119- 183. Cichan. M. A.. and T. N. Tay lor. 1982. wood-borings in Arnold, C. A. 1945. Silicified plant remain s from the Me­ Promnoxvtnn: plan t-animal interactions in the Ca rbonif­ sozoic and Tertiary of western North America . I. Ferns. erous. Palaeogeogr. Palaeoclirnatol. Palaeoeool. 39:123­ Michigan Acad . Sci.. Arts leiters Pap. 1944. 30:3-34. 127. --. 1958. A new Tempsk)'Q_ Contrib. Mus. Pakontol. Cohen, A. W., W. Spackm an . and R. Raymond. Jr. 1987, Univ. Michigan 14(8):133-142. Inlerpreting the: cha racterist ics ofcoal sea ms from chemi­ Ash, S. R.• and C. B. Read . 1976. North America n species cal. physical. and petrographic studiesof peal ~IS . Pages of Tt'mpskyaand lheir stratigraphic significance. U.S. Gcot. 107- 125 i n A. C. Soon. 00. Coal and coal-bean ng strata. Surv. Prof. Pap. 847:1---4 2. Recent Advances. Geol ogical Society Special Publication Bonefante-Pasolo. P. 1984_ Anatomy and morphology of no. 32. Geological Society ofAmerica, Boulder. Colo. VA mycorrhizae. Pages S--33 i n C. L Po.... ell and O. J, Corda. A . J. C. 1845. Beitrage zur Flora der v orw eh. J. G. Bagyaraj. eds. VA mycorrhiza . CRC Press. Boca Raton, Calv'sche, Prague. (See p. 87. pl. 47, fig. 7.) Aa. Crabtree, D. R. 1987, Angiosperms of tbe nonhern Rock y Boodle, LA. 189 5. Sporn in a specimen o f Tt'mpsk}7J Moumains: Albian 10 Campanian (Cre taceous) megafossil (EnJOKf'nites). Ann. Bot. 9:13 7-141. 1I0faS, Ann. Mo. Bot. Gard . 74:707- 747. Brown. R. W. 1950. Cretaceous ptants from southwestern Crebe r. G. T. 1977. Tree rings: a nat ural data-sto rage sys­ Colorado. U.S. Geol, Surv . Prof. Pa p. 22 1-0 :45-66. tern. Bioi. Rev. (Lond .) 52:349-383. Casagrande, D. J. 1987. Sulphur in peal and coal. Pages 87­ Creber. G. T., and W. G . Chalcner. 1985. Climati c ind i­ 105 i n A. C. Soon. 00. Coa l and coal-bearing strata. Recent cations from growt h rings in fossil woods. Pages 49- 74 in Advances. Geo logical Societ y Special Publicatio n no. 32. P. 1. Benchley, ed. Fossils and clim ate. Special issue of the Geo logical Society of America, Boulder. Colo. G('% gical Journal. Wiley, New York. 528 I NTER NA TIO NAL JOU RNAL O F PL ANTSC IE NCES

Cross. A. T..E. B. Max field . E. Cotter, and C. C. Cross . 1975. Seward. A. C. 1899. On the structure and affinities of .'.la ­ Field guide and road log to the western Book Cliffs. Castle tonia peairuua R. Br., with notes on the geological history Valley and pans of the Wasa tch Plateau . Brigham Young of the Matonia ccae. Philos. Trans. R. Soc. Lond. B 191: Univ. Geol , Stud. 22(2): 1- 132 (See p. 173- 174.) 171- 209. Hall, J. R. 1984. o(VA mycorrhiza l fungi. Pages ---. 1923. The use cruie microscope in palaeobotanical 57- 94 in C. L Powell and D. J. Bagyaraj. eds. VA myco r­ research. J. R. Microsc. Soc. 43:299- 302. rhiza. eRe Press. Boca Raton, Fla. --- . 1924. On a ncw speciesofTt'mpskya from M ontana. Hdntzchel. W.• F. E1 -Baz. and G . C. Amstutz. 1968. Co p­ Tl!mpsk)'tl kno...-honii. sp. nov. Ann. Bot. 38:485-507. robtes, an annotated bibliography. Mem . Oeot. Soc. Am. Spackman, W.• Jr. 1948. A dicotyledonous wood found as­ 108:1-132. scctated with the Idaho tcmpskyas. Ann. M o. Bot. Gard. Harley, J. L . and S. E. Smith. 1983 . Mycorrh izal s)"mbiosis. 35:107- 116. Academic Press, New York. Stach. E.. M. Mackowsky, M. 'reicbmoner. G. H. Taylor, D. Hlu stik . A. 1990. Bohe mian biographies III: the story of Chandra, and R. 'rek hmuner. 1982. Stach's tu.tbook of r emp$kya. lOP Newsl. 42:12-13. coal pelrology, 2d cd. Gebnjder Borntraeger, Berlin. 535 Jackson, R. M.. and P. A. Mason. 1984. Myoorrilil'.a. Ed­ ward Arnold. Baltimo re. Stokes""., C.. and P. B. Webb. 1824. Dcscnprions of some Katich. P. J., Jr. 1951. Recent evidence for Lower Creta­ fossil vegetables oflhe Tjfgate Forest in Sussex . Goof. Soc. ceous deposits in Colorado Plateau. Am. Assoc. Pel. Geot. london Trans.• 2d scr., 1:423-426. Bull. 35( 19):2093-2094. Sto kes, W. L. 1952. Lower Cretaceous in Colorado Plateau. ---. 1952. Occurrence of Tempskya in the Lower Cre­ Am. A~ . Pet. Geo!. Bull. 36:1766-1776. taceous of the w estern Interior (Utah). 1. Paleonto1. 26: Th ayn. G. F" and W. D. Tidwell. 1984. fl ora ofthe lower 677. Cretaceous Cedar Mountain Form at ion of Utah and Col­ McKnighl. K. II.. and J. K. Rigby. 1963. Lower Cretaceous orado. II . Mesemhnoxylon nokrsi. Great Basin Nat. 44: Temmkya from eastern Utah (abstract). Geol. Soc. Am. 257-262. Spec . Pap. 73:91. Thayn. G. F.. w. D. Tidwell.and W. L. Stokes. 1983. flora Nelson, M. Eo, and D_M. Crooks. 1997. Stratigraphy and of the Lower Cretaceous Cedar Mountain Formation of pajeomclogy of the Cedar Mountain Formation (Lower Utah and Colorado. I. Paraphyllanthoxylon utahense. Great Cretaceous), eastern Emery County, Utah. PagC'S 55-63 in Basin Nat. 43:294-402. W. R. Averell , ed. Paleontology and geology of the Dino-­ - - -. 1985. Flora of the Lower Cretaceous Cedar Moun­ saur Tria ngle. G uidebook 1987. Museum w estern Colo-­ lain Formation of Utah and Colorado. III. 1