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PRINCIPES IVol. lI FossilPalms

JosepnA. Ture, M. D.

Living palms are familiar to oryas (4, p. 172) mentionsthat the dis- "one all who grow them and are a character- covery was of the most significant istic feature of many parts of the trop- finds in recent times" although with in- ics. That palms have a long history in complete evidence it should be classi- time is not always recognized.It is the fied only as palm-like. Ladd and Brown object of this article to illustrate some also illustrated in color the cross-sec- o{ the palms found in the United tion of a fossil palm trunk of a later States, correlating structure of stems, era found on Antigua in the West roots, and leaves with those seen in Indies. It has a light yellow-brown color living palms, the anatomy of which has similar to much of the fossil palm wood beenreviewed and evaluatedby Tomlin- found in Southern California. son in his volume on Palmae in Anato- Arnold (7, p. 341) placesmost of the my of the (L). fossil palms in North America along Palms first definitely appearedin the with rocks of the epoch (36'58 Upper rocks, some 63-90 million years ago). There was a shi{t' million years ago by the geologic time ing or migration of palms southward scaleof Kulp (2). A fossil which has due to climatic changes at the end of been called Palmoxylon clillwoodensis the Eocene which continued into the dating from that period has been found Pliocene(I-13 million yearsago). Thus in the Magothy formation of New Jer- the occunence of palms in the fossil sey wheretoday no palm lives (3). De- record has served more or less as an Ievoryas (4, p. 173) mentions that indicator of the prevailing climate of palm-like leaf impressionsfrom France the past. No6 (B), La Motte (9) and (the Propalmophyllum of Lignier) were Mahabal6 (10) give many references reported in 1907 as dating from the Ju- on {ossil palms. rassic period ( 135-IBI million years There are several imPortant areas ago). If truly a palm, this stands as {rom which material of fossil palms has evidencefor the presenceof flowering been obtained. One of the most fam' plants at that early date. However, go- ous is the extensive region along the ing back still further, the discovery of coastal plain of the United States bor' a palm-like leaf imprint in sandstone dering the , where rocks was reported and illustrated in color containing what is known as the Wil- (5) by Ladd and Brown and by Brown cox flora were laid down in lower (6) Triassicperiod (181-230 from the Eocene times. These beds contained A million years ago). This ancient fossil many including Chamaedorea' leaf was found by Dr. G. Edward Lewis lt like species.At about the same time, near Placerville,Colorado, in the vicin- existedin northern New ity of the San Miguel River. Several the Raton flora months later, Dr. Brown and Dr. Lewis Mexico.and southernColorado. Knowl' explored the area where the first im- ton (11, p. IB0) includesa photograph print had been found. Six more im- of a leaf impressiono{ a Sobal-likepalm prints were uncovered and taken back from this flora. From a succeeding to the Smithsonian Institution. Delev- middle Eocene flora, the Claiborne r9671 TUTA: FOSSIL PALMS 55 which extendedfrom Alabamato , in time, from the upper Eocene Berry (12, p. 5I) described several Jackson flora of Texas, fossil fruits o{ palms basedon leaf structure-one sug- a date-like palm were found. Chaney gestive ol Thrinax, and two feather- (13, p. 1I) publisheda photographof leaved speciescalled Baetrites and.Geo- a large bed composedof layers of fossil nomites from their resemblanceto mod- palm leaves from the Clarno shale of ern Bactris and, Geonoma. Still later Oregon, also of Eoceneage. Most of GElLOGICTIME SCATE 'j;:"[H:t"(') PERIOD POSITION OF FOSSIL PALMS bEGINNING & DTJR,ATION IN CEOIOGIC TIME SCALE ;ND OF PERIOD *PALMS IN AUITIORIS COLLECTION RECENT 0 - 0.01 0.01. I lo onn u PLEISIIOCENE- a 0.0r.- r.ot L aER(), TURIDA PLIOCENE IRICARDO BEDS upPer 1-13 L2 (I.AST CTIANCECANYON) CALIF(N,NIA :oPIAH COI'NTY, MIS$SSIPPI IROSAIT{OND,CALIF. (NEAR MOHAVE) IAPAIT (TERTIARY) EPOCH NOT KNOL'N uPper IBARSTOC BEDS (MULE CANYON) CALIF. . mlddle L3-25 L2 KTEHACHAPI, CALIF. (HORSE CANYON) lower TIAGRANGE, L FAYETTE COUNTY, TEXAS 25-36 t1 INTIGUA, WEST'INDIES ? OLIGOCENE fr SUB-DTVISTON NOT I(NOT.'N EOCENE 36-58 22 TFARSON, [tYO,tINc (EDEN-VALLEY) ll TNEPHI,, H UTAH : kSIERRA COI'NTY, NEtr MEXICO E- upPer io-+s 9 :LARNO, OREGON, FOSSIL LEAVES 01 IACKSON.FLORA - GULF COAST REGION ]RAZOS CO. TEXAS. NIPA.LIKE FRUITS T{ nlddle 45-52 7 - E{ :I,AIBORNE.FLORA GULF COAST REGION TATON.FIORA NORTHERNNEW MEXICO lower 52-58 6 IILCOX FLORA GI'LF COAST REGION IIPA-LIKE FRUITS GRANADACO. MISS. WORTH CAROLINA PALEOCENE ,7 -63 5 IRAZIL. NIPA-LIKE FOSSIL FRUITS

CRETACEOUS OSSIL PAII{S FIRST APPEAR IN UPPER-li..l uPPer 63-90 27 LEI. _IERIgD (MAcoruy rmuruoH) Iower 90 - 1.35 45 IT'RONIAN OF FNAN.:N JI'RASSIC - r35 - 181. 46 ,AW-LIKE I.EAF IMPRINTS N,OPALMOPHYLLIJIT{OF LICNIER (FRAIICE) LEl - 230 49 IAIX.LIKE LEAF IMPRINTS (COIORADO'-

PER},TIAN l3o - zao 50 PENNSYLVANIAN 180 - 320 t8 MISSISSIPPIAN t20 - 345 25 Jl ,45 - t+Os 60 lO5 - 425 20 NO EVIDENCE OF pAL!,rS u ;25 - 500 75 OR CAI{BRIAN '00 - 500 100 PAI.Iq.LIKE STRUCTT'RES

INCLUDES THE BEIOII THE TRIASSIC PERIOD OLDEST BASEMENT ROCKS FROM THE 600 - 3600+ 3000+ TIME OF M,IGIN OF THE EARTH UP TO THE LOWERCAI{BRIA PERIOD ! PRINCIPES lVol. 1r the {ossil palms b{ Wyoming occur in Eocene age have also been found in rocks o{ the same age. Belgium, Borneo, France, India and Fossil palms in Texas are found in Russia. Some of these structures rocks from both the Miocene and have also been found in beds of Eocene epochs, those from Fayette Miocene age in India. Fossil pollen County being of Miocene age. In Cali- grains have beenreported frorn Borneo fornia, the Tehachapi {lora of the pres- in beds of Cretaceousage. In the Uni' ent western Mohave Desert region was ted States Nypa-like {ossil fruits have laid down in the Middle Miocene(about been{ound (Berry 19. p. 176) in the 18 million yearsago) while the Barstow Wilcox (Eocene)flora, Granadaforma- beds, which include the Mule Canyon tion, Granada Count-y, Mississippi. t i area north of Yermo, are of upper Mio- From Texas Berry (12, p. 150) re- q ceneage (I5 million yearsago) and the ported similar fruits found in the Eo- Ricardo beds of Southern California, cene Fayette{onnation near Wellborn. which occur in the Last ChanceCanyon Brazos County. Also {rom the Paleo- area, are o{ lower Plioceneage. Axelrod cene of Brazil. {o,ssilfruits oI lYypa (14) describedfossil palms of Plio- havebeen reported by Dolianiti (20 ) in cend age from Palmdale,California. An 1955. These structures can dri{t {or imprint of a Sobal-likeleaf with a costa- long distanceswith the oceancurrents. palmate blade was revealed during the The oceanicpaths of migration are dis- processo{ road building near Castaicin cussedby Corner (21, pp. 249-252\. Los Angeles County and is illustrated He also suggeststhe possibilit,vthat the by Hertrich (15, p.4). From Japanon .Nypo- like fruits found in southenr the islands of Hokkaido and Kyushu. U.S.A. may be from tropical Anrerican Kryshtofovich (16) has describedSa- palms other than l{ypa. bal-like palm leaf impressions. In Eu- One of the most inforrnative de- rope, during the Focene epoch, Sabal- scriptionsof how plants becarrefossils like palms grew in England. France and is givenby Arnold (7. p. 14-40).Com- southern Russia and also appear in the pressions,casts and petrifications(per- Pliocene rocks of the Rhone Valley. mineralization)are the most inrportant' Eocenebeds of England occur in two methods by which fossilization is e{- separatebasins, the Hampshire and the fected.O{ the monocotyledonousplant London. Fossil fruits o{ the tropical forms, the petri{ied (silicifiedI palnrs palm Nypa (Nipa) have been found in are among the best preserved.Manl' the old delta mud or clay which lies be- factors are involvedin the petrification low London. It is also exposedin the process.One importantfactor for sel- clif{s of eastern Kent County and the ting the stage for petrification is the Isle of Sheppeylocated at the mouth of rapid submergenceof the trunk and the Thames River. Reid and Chandler roots in a body of water where ox1'gen (17) and Chandler(18) have made ex- is absent. Another important factor is tensive studies on the flora of the Lon- deposition in the water of finely di- don clay in which fossil /Vypo struc- vided sedimendssuch as clay. nrud" tures have been ,found. Geologically sand and volcanicash. this important palm has existed since The complexpetrification process has the Cretaceousperiod. At the present been studiedby Arnold (22), Darrah time lVypa is restricted to parts of (23) and Barghoorn (24, 25). Ther' southeastAsia and some of the South have produced much evidenceindicat- Pacific Islands. Fossil Nvpa fruits o{ ing that the {undamentalbasic process TUTA: FOSSIL PALNIS

1. Fossil palm, Tehachapi,Kern Co., Calif. Photograph enlarged3 X. Trro x-vlemvessels- occasionallythree, in each vascularbundle stand out like ink dots. \\'ith the use o{ a 4X or 10X lens some bundlesshow groups of phloem structuresbetleen the r,vlem vesselsand the "cap", large fibrous bundle sheath also .agatizedsepta diliding the palm structure into lslan d s.

2' Fossilpalm' Horse can;'on'"5"";:f:;,i:f'j";ili;" sr'orringthe intricate |:l?,'J.'iLiioiili'"" is chiefly one of infiltration. rather than suhstances. The nature of the process "molecule the by molecule" replace- can be shown in preparations using a ment o{ the structure b1. mineral highl,v carbonaceous black Wvonrin6q PRINCIPES lVol. 11

3. Fossil root mass, Mule Canyon, Yermo, California. The central area at (A) devoid of root structures, has typical vascular bundles and represents the lowermost portion of the stem, The opposite or upper side of the specimen shows stem structure with vascular bundles and a narrow nm showlng roots.

fossil palm. Microtome sections and the study of detailed structure, one of "peel" prepbrations were made follow- the incentives for collecting petrified ing the techniquesdescribed by Darrah wood (chiefly stems and roots) is the (26), Joy, Willis and Lacey (27) and wide variation in color and pattern in Kummel and Raup (28). The silica was the polished sections. Most palm wood removedwith hydrofluoric acid, leaving is a yellow brown, some is multicolored a residual organic carbonaceousframe- with red, black, green and blue, in fact work which outlines the cellular pattern every imaginable color combination. of the fossil structure. This material In general,however, palm woodsare not representsthe degradation products of as colorful as the diffusely agatized cellulose, hemi-celluloseand lignin, of silicified Araucarioxylon arizonicum oI which the original woody structure o{ the Petrified Forest National Monument the plant cell walls is chie{ly composed. of Arizona, or the Utah Cycadeoids. The complex structure of lignin has Certain of the fossil palms show excep- recently been reviewedby Brown (29). tionally well preservedvascular bundle Figures 15 and 16 compare the peel and . Naumann (30) has preparation with a standard thin sec- described the variation in color found tion, both methods showing a similar in palm wood from Fayette County, carbonaceousframework. Texas, and especially has given a de- tailed account of searching for fossil Collecting Fossils palms in that region. Apart from providing material for Only a few specimenshave been il- re67I TUTA: FOSSIL PALMS 59

t?:::..:.,1 :;:,;i::)ii 4. Semi-castof black Wyoming fossil palm shows trunk rings (leaf base scars) in the sur{ace layers of the stem, also vertical fissures in the spacesbetween the rings. The interior consists of sandstone and black areas containing crystallized and carbonaceous remnants of the vascular bundles and grouno Itssue.

'rvith 5. Wyoming fossil palm spiny stems shows numerous basesof spines which had been broken off close to the stem either before or after petrification. At (A) the stem structure had been disrupted by some severemechanical force before petrification had taken place. 60 PRINCIPES lVol. 1l lustrated in this article to demonstrate roots. The junction of the inner surface the variety of fossilsone may encounter. of the adventitious root mass and the Choice specimensare not easy to find cortex of the stem shows some of the today since most material near the roots lying at right angles to the cor- surface has already been taken by pri- ter and entering the ground tissue for vate collectors. Most of the material a distance of about a millimeter. In has been collectedin Southern Califor- two specimens the adventitious roots .Wyoming. nia, Utah and Other loca- were unequally distributed. some areas tions from which fossil palm specimens having a layer of matted roots up to I have been obtained are Arizona, Baja inch in thickness,whereas on adjacent California (Mexico), ,Missi- areas of the same specimen,the roots ssippi, southern Nevada, North Caro- do not form a thick layer but show lina, southwest Oklahoma and Texas. single closely spaced short spine-like The Arizona fossil wood came from a structures perpendicular to the stem. mine 50 miles north of Wickenburg. In some places, two or three of these and contained visible yellow areas o{ spinesare conjoined and cappedby a radioactivematerial, probably carnotite. bark-like silicified structure. The spines The specimensfrom Searchlight, Neva- show a root structure and their appear- da have been thought by some to be ance is somewhatsuggestive of the root- petrified Joshua Tree. Wyoming has spines seen on stems of living Cryoso- furnished the greatest variety of fossil phila. palm material, most of it being in rocks Cane.like Stems of Eocene age from the Eden Valley Many fossil palm stemsare thick but area near Farson, Wyoming. The ma- someare slenderand cane-like,resembl- terial includes stems with bark-like ing those o{ such modern genera as outer structuresshowing leaf base rings Chrysalid,ocarpusor Geonoma.In Wyo- and longitudinal fissures, adventitious ming, cane-like fossils are found in a roots, branch-like structures, cane-like small valley about one mile across at stems, spiny stems, underground root its greatestwidth in Sublette Co., near massesand specimenswhich are sug- the Big Sandy Reservoir north of the gestiveof damageby insectsand fungi. town of Farson. The floor of the val- Some examples o{ these structures fol- ley is coveredwith sage brush and sur- low. rounded by rolling hills about 300 feet high. The matrix which held some of Fossil Palms with Adventitious Roots the stemsappears to be composedchief- Sorne of the Wyoming stems show ly of limestonewhich dissolvesin hydro- an overlying layer of adventitiousroots chloric acid leavinga small silicateresi- up to Ll2 inchesin thickness(Fig. 7). due solublein hydrofluoric acid. Most The outer surface of the adventitious of the cane-likespecimens are found root mass is coveredwith a bark-like free from their matrix. layer up to 0.15 inches in thickness. One of the external molds where the In the outer layer, somespecimens show stem had fallen out o{ the matrix shows rings o{ Ieaf scar origin. well-defined plainly the outlineso{ rings (leaf base It is interestingthat this outermostlayer scars) on the trunk (Fig. l0). A few covering a thick layer of roots resem- specimensare black due to a high con- bles closelythe bark-like outer surface tent of carbonaceousmaterial. The layers of many o{ the Wyoming speci- stemsare mostly short sectionsthough mens which do not have adventitious someare up to 12 inchesin length and 1967I TUTA: FOSSILPALMS

6' wvomingrossil palm spinesscatt€red through' :ilndltil;::T:,'JJ;',f,:T#i'.,'1.".:t

7. Wyoming fossil palm with thick zone of adventitious roots. (A) cortical zone of stem. (B) central zone of stem agatized in a variegated fashion. (C) zone of adventitious roots. (D) depositsof fossil algae. 62 PRINCIPES fVol. 11

B. Wyoming black fossil palm with central area of white agate formation. The stem is covered by a thick layer of adventitious roots, also an overlying bark-like surface layer showing trunk rings. The spaces between the rings show a gray to white surface layer marked by roughly parallel ver- tical fissures. \r/2 inches in diameter. An occasional trunks. stem with attachedroots has been found Spiny Stems When the ends are polished some of Severalexamples of stemswith spiny them show glistening quartz crystals structures are found among Wyoming lining cavities; others have jet black fossil palms. The spines have been areasand foci of blue agate.The broken broken off close to the stem due to ends are roughly perpendicular to the mechanicalforces which occurred be- long axis. Why they break up or frac- fore or after the processof petrification ture in particular manner is not this 'Visitors had taken place. Only scarsof the bases easily explained. have asked of the spines or short stumps remain. this question of the park naturalists in The spines are scattered diffusely the Petrified Forest National Monument throughout the internodes between the of Arizona (3I). They see the huge trunk rings (Fig. 6) without a definite petrified logs with the ends looking as distribution pattern, exceptin one speci- though they had been sawed by man, men where the spines were lined up in instead of having been broken by the a more or less parallel fashion. No fos- mechanical forces of nature. One ex- sil palms were {ound where the spines planation given is that shock waves were lined up along the lea{ scar rings. causedby earthquakesinitiate rhythmic vibrations which caused more or less Damage from Insecls and Fungi regularly spaced breaking up of the A distorted specimen of fossil palm 1967l TUTA: FOSSILPALMS 63

9. Wyoming fossil palms rsith cane,like stems. The spaces between trunk rings vary {rom 2-10 cm. in length. The sur{ace layers have partially disappeared most o{ the speci- mens probably due to weathering.

l*""*:*:t'f.:1m:*[:"[l;'lll"-,i,ilii;"|,f:ll:'[;i:l:.Xfl3;;^#:",iflili ;f::,,T;"h:"#; an external mold with trunk ring markings. [\/ol. ll 64 PRINCIPES

Polishedcross- ll. Distortedfossil palm (diamererof stem18 cm.), Mule Canyon,Yermo, Cali{. airirr.ri*[;i;; ;.;y ;r+.g wiih a {er- areas having recognizable.vascular .""ri--.rr""ilg-* be{orepetrification' [r"aj".."i" ,-frJ"i"u ," ifr"i"fi-;;;;t;;"i tiny holessuggesti'e of beetleborings Peelpreparations and thin sectionsshowevidence of fungusinYol\ement'

:.r"iJ,:. j

end o{ trunk (X) and root 12. Fossil palm stump from LaGrange' Texas-'-Junction o{ lower measurinq22 X lB X 17 inches and weighing 230 lbs. 1967l TUTA: FOSSIL PALMS

"::ta,aa::::,:::'i:,.a:t,,,:.:,',..,::, .,1:,:,,,\ ..;., :: Fossilpalm leaf imprint.

14' Fossilpalm leaf imnrint, HorseCanyo-n, Kern Co.,Calif. There is a continuationof the petiole as a rib extindinginto ("o.iuiul-",").'I. ihe bi"d" irr" p"iloi" *;ttr''li. .]r,"'".o""sion,rhere is an areawhich is unfortunatelymissing. rl 66 PRINCIPES [Vol.

peel preparation' lS. Black Wyoming fossil -palm (diameter of stem 2.5 cm.). Photomicrograph of masnification30x, enlarged2.5X.

(Fig. 11) from Mule Canyon north of segmentsof the circumferenceof some Yermo, California is of interest in that of the lamellae. They are not visible cellulose-acetatepeel preparations with polarized light. Also scattered showed filament-like structures sugges- throughout some of the micro'crystal- tive of fungal involvement.Also on the line material are single or clumped fila- surfaceof one end of the samespecimen ment-like structures. Fine fracture lines are several small superficial borings are observed in the micro-crystalline possibly due to insect damage. These material. A thin sectionfrom this speci' might conceivably have served as the men was sent to Dr. E. S. Barghoorn point of entry for fungi, but there is no of Harvard University who agreed that way to prove such a relationshiP. the filamentous structureswere of fung- Stevens (31) has reported Several thin sections (30 microns al origin. fungal hyphae in the central thick) weremade from the mottled gray finding and roots of a fossil palm com' areasof the specimenshown in Fig. 11. cylinder the uPPer Cretaceous,Mon- Most of these thin sections show fila- ing from Sea Bright, New Jer' ment-like structures. Areas are also mouth {ormation, seen containing severalconcentric lam' sey. ellar bodies which with ordinary light Acknowledgemenls been show filament-like structures occupying This project could not have 1967I TUTA: FOSSIL PALMS

16. PhotonricrographoI thin section.fromsame palm a,s15, magni{ication ':;;;;-'iln-0."i*yr"-.JgX, enlarged2.5X. PH-phloem.XY-xvjem vessets.FB-fib.*. r""jr"' .h"";h ;;' GR-racunae rn groundttssue.

carried out without the help of inter- kie Young, hospital librarians, obtained ested friends who gave of their tine referencesthrough the inter-library loan and experienceand valuable specimen arrangement.Mr. and Mrs. Leo D. Bern- material. I am indebted to Dr. Chester er of Glendora,Calif. and Mr. and Mrs. A. Arnold, Professor of Paleobotanyat J. D. Rittenhouseof Azusa" Calif. have the University of Michigan for his gen- beautiful and valuable collections of erous help in identifying many fossil petrified material which I have had the plants from the western states. Dr. E. opportunity to study. Mr. and Mrs. S. Barghoorn kindly examined one of Harry F. Ohlsen of Wilmington, Calif. the fossil palm specimensfor fungal have furnished severalof the Wyoming involvement. Mr. and Mrs. Samuel E. specimens,which they collected over a Kirkby of Riverside, California have period of 25 years. Their exceptional been generousin making their private interest in unusual petrified material collection of plant fossils available for has beenparticularly helpful. Mr. James study, and also for giving expert ad- Frank Britsch of Long Beach, Calif. vice on fossil material. Dr. H. H. Dib- A. Greene,Mr. Archie Greeneand Mr. bern, Dr. M. Itano and Dr. J. W. Rev- have been of great assistancein cutting nolds, associatesin the Dept. and Patir- and polishing many of the specimensl ology, Memorial Hospital of Long Mr. and Mrs. Lowell Gordon. Long Beach, Calif. and Dr. Ruth Russell of Beach Gem and Mineral Dealers. have the Long Beach State College have furnished me with severalMule Canvon. helped with much of the photographic Yermo, Calif. specimens. Dr. Grant work. Miss Francis Ishii and Mrs. Jac. Beckstrand of Palos Verdes Estates, 6B PRINCIPES [Vol. 1l

17 & fB. Wyoming' black fossil palm, diameter of stem 5 cm. Microplotographs using polaroid film, Leitz-Wetzlar metallograph using polarized light'

17. Magnification Xl00 taken from peripheral portion of central cyl]nder shoning a "ingle t'capr'. vascular"bundle. FB-fibrous bundle pfl-phloem structures. XY-two xylem \.essels. PR-protoxylem.

Calif., palm enthusiast,gave much en' ,), Berry, E. W. 1916. A petrified couragement and valuable suggestions. oalm from the Cretaceousof New Members of the Palm Society of South- 1".."y. American Journal of Sci- ern California who have beenmost help- ence ser. 4, 4l (whole number ful and generouswith time in giving in- 19l): I93-197. formation on living palms are Mr. Ho' 4. Delevoryas, T. 1962. Morphology race Anderson of Leucadia. Calif., Mr. and evolution of fossil plants. Holt, David Barry, Jr., of Brentwood, Calif., Rinehart and Winston, N. Y. Mr. Otto Martens of the Diegaard Nurs- 5. Ladd, H. S. & R. W. Brown. 1956. eries in Monrovia, Calif., Mr. F. A. Tet- Fossils lift the veil of time. Na- ley II and Mr. F. A. Tetley III of Co- tional GeographicMagazine 109: rona, Calif. and Mr. L. M. Whitelock of 363-386. Eagle Rock, Calif. 6. Brown, R. W. 1956. Palmlike plants Literature Cited from the Dolores formation (Tri' I. Tomlinson, P. B. 1961. Anatomy assic) southwesternColorado. U.S. of the monocotyledonsII. Palmae, Geological Survey Professional C.R.Metcalfe,ed. ClarendonPress, Paper 274-H: 205-209. Oxford. 7. Arnold, C. A. 1947. An introduc- 2. Kulp, J. L. 196I. Geologic time tion to paleobotany.McGraw'Hill, scale.Science 133: I105-1114. N. Y. 1967l TUTA: FOSSIL PALMS

lB. Magnification X50. Upper zone: cortex. Lower: peripheral zone of central cylinder with intricate ramifications oI leaf base complexes.

o o. No6, A. C. 1936. Fossil palms, in 13.Chaney, R. W. L948 (reprinted Dahlgren, B. E. Index of American 1956). The ancientforests o{ Ore- palms. Field Museum of Natural gon. Condon Lectures. Oregon History, Botanical Series,14: 44I- State System of Higher Education. 456. Eugene,Oregon, 9. La Motte, R. S. 1952. Catalogueof t4. Axelrod, D. I. f950. The Anaverde the Cenozoicplants of North Amer- flora of Southern California. Car- ica through 1950. The Geological negie Institute of Washington Pub- Society of America, Memoir 5l: lication 590: I44, pl. 2, l. 2. 239-242. Also see under specific 15. Hertrich, W. 195I. Palms and cy- palm genera. cads.Henry E. Huntington Library 10. Mahabal6, T. S. 1959. Resolution and Art Gallery, San Marino, Cali- of the artificial palm gents Palm- fornia. orcylon: a new approach. Paleo- 16. Kryshtofovich, A. I9IB. Occurrence botanist 7: 76-84. of the palm, Sabal nipponica, n. I1. Knowlton, F. H. 1927. Plants of sp. in the Tertiary rocks of Hokkai- the past. A popular account of fos- do and Kyushu. Journal of the sil plants. Princeton University Geological Society of Tokyo 25: Press, 59-66,fig. 1,2, 3. 12. Berry, E. W. L924. The middle and 17. Reid, E. M. and M. E. J. Chandler. upper Eocene floras of southeast- 1933. The London Clay Flora. ern North America. U. S. Geolog- British Museum(Natural History), ical Survey Professional Paper 92. London. 70 PRINCIPBS [Vol. ll IB. Chandler, M. E. J. 1964. The low- 24. Barghoorn, E. S. f952. Degrada' er Tertiary floras of southern Eng- tion of plant tissues in organic land IV. British Museum (Natural sediments.Journal of Sedimentary History), London. Petrology 22:34-4I. I9. Berry, E. S/. fqf6. The Lower Eo- 25. Barghoorn, E. S. 1952. Degrada- cene floras of southeasternNorth tion of plant materials and its rela- America. U. S. Geological Survey tion to the origin of coal. Second Professional Paper 9I. conferenceon the origin and con- 20. Dolianiti, E. 1955. Fructos de Nipa stitution of coal. Crystal Cliffs, no Paleocenode Pernambuco,Bra' Nova Scotia.lBI-203. de Agricul- sil. Boletin Ministeria 26. Darrah, W. C. 1960. Principles of 1'36. pls. tura Rio de Janeiro 158: Paleobotany, ed. 2. Ronald Press, t2 N. Y. 2r.Corner, E. J. H. 1966. The Na' K. W., Willis, A. and W. S. tural History of Palms. University 27. Joy, J. A rapid cellulosepeel of California Press.Berkeley, Calif. Lacey.1956. in paleobotany.Annals 22. Arnold, C. A. f941. The petrifica' technique Botany,ser.2,2U 635'637. tion of wood. The Mineralogist 9: of 323.324,353-355. 28. Kummel, B. and D. Raup. 1965' 23. Darrah, W. C. L94L. Changing Handbookof paleontologicaltech- views of petrification. Pan Amer- niques.W. H. FreemanCo., San ican Geologist76: 13-26. Francisco,Calif.

I. Allagoptera arenaria cultivated at Fairchild_Tr_opical Garden with an apparently dichotomous crown. Photo M. V. ParthasarathY. 1967I TUTA: FOSSILPALMS 7T 29. Brown, S. A. f966. Lignins. An- fied Forest Museum Association. nual Revue of Plant Physiology Holbrook. Arizona and the Arizo- L7:223-2M. na State Highway Dept. 30. Naumann, R. C. 1964. Wood re- 32. Stevens,N. E. 1912.A palm from placementsof Fayette Co., Texas. the upper Cretaceousof New Jer- The Lapidary Journal IB: lB7-I91. sey. American Journal of Science 31. Brodrick, H. J. I95I. Agatized ser. 4, 34 (whole number 184) : Rainbows. Popular Series 3. Petri' 42T-436.

I I I I

2. Close-up of plant shown in Fig. l. Developing leaf primordia belonging to two buds areindicated by arrows.'Theleaf tffi"$]"FjL*T.:,il: ".o*" is held by the hand. Photo