A Flora from the Dakota Sandstone Formation (Cenomanian) Near Westwater, Grand County, Utah

Brigham Young University Science Bulletin, Biological Series

Volume 14 Number 3 Article 1

9-1971

A flora from the Dakota sandstone formation (Cenomanian) near Westwater, Grand County, Utah

Samuel R. Rushforth Department of Botany and Range Science, Brigham Young University, Provo, Utah

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OCT 29 1971 Brigham Young University HARVARD UNlVERSlBCi Science Bulletin

A FLORA FROM THE DAKOTA SANDSTONE FORMATION (CENOMANIAN) NEAR WESTWATER, GRAND COUNTY, UTAH

Samuel R. Rushforth

BIOLOGICAL SERIES — VOLUME XIV, NUMBER 3

SEPTEMBER 1971 BRIGHAM YOUNG UNIVERSITY SCIENCE BULLETIN BIOLOGICAL SERIES

Editor: Stanley L. Welsh, Department ot Botany, Brigham Young University, Provo, Utah

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A FLORA FROM THE DAKOTA SANDSTONE FORMATION (CENOMANIAN) NEAR WESTWATER, GRAND COUNTY, UTAH

Samuel R. Rushforth

BIOLOGICAL SERIES — VOLUME XIV, NUMBER 3

SEPTEMBER 1971 TABLE OF CONTENTS

Page

INTRODUCTION 1

GEOLOGICAL BACKGROUND 1

Litliological Cliaracteristics 1 Geology 4 THE AGE OF THE DAKOTA SANDSTONE 4 A CONCEPT OF A DAKOTA FLORA 7 PALEOECOLOGY 9 PALEOCLIMATE 12 SYSTEMATICS 13 Genus Equisctum 13 Genus Asplenium 14 Genus Conioplem 18 Genus Ildusmannia 18 Genus Gleichcnin 20 Genus Matouidium 27 Genus Astrulopteris 34 Genus Cliulophlehis 34 Genus Ilex 38 Genus Maniioliu 38 Genus Ficus 38 Genus Eucali/ptus 40 Genus Platanus 42 Genus Salix 42 ACKNOWLEDGEMENTS 42 REFERENCES CITED 43 A FLORA FROM THE DAKOTA SANDSTONE FORMATION (CENOMANIAN) NEAR WESTWATER, GRAND COUNTY, UTAH

Samuel R. Rushforth'

ABSTRACT

A Cretaceous (Cenomanian) flora from the coloradica represent the dominant forms in this Dakota Sandstone Fomiation near Westwater, flora. Grand County, Utah contains an admixture of The Westwater flora contains fourteen genera ferns and angiospenns. The ferns of this flora including nineteen species and one variety. New are representative of an older Jurassic-Wealden species described from this flora include Asplen- \egetational type, whereas the angiosperms are ium dakotensis, Coniopteris westwaterensis and t\pical of the modem vegetational type. Species Ilex serrata. of GleichenUi and Matonidhim and Astralopteris

INTRODUCTION

A fern-angiospenm flora from the Cretaceous distribution patterns of manv previously de- (Cenomanian) Dakota Sandstone Formation has scribed species. The Westwater flora is one of been under studv for some time. This flora was few paleofloras which illustrates an admixture collected from an ash seam and sandstone in of an older Jurassic-Wealden floristic tvpe with the Dakota Sandstone Fonnation from Rabbit a modern angiospermous floral tvpe. Third, \'alle\', Grand Countv, Utah, and along the this flora provides new information on a time road and surrounding areas between U.S. High- of the earth's history when angiosperms were wav 50 and Westwater, Grand County, Utah. expanding from a position of little floristic im-

This Dakota flora is significant for three portance to a position of dominance in Cre- reasons. First, it contains several new species taceous and later floras. of fossil plants. Second, it extends the known

GEOLOGICAL BACKGROUND

LiTHOLOGic.\L Characteristics.— Two early Lesquereux (1874) presented an excellent workers, Marcou (1864) and Capellini (Capel- discussion on the probable origin of the Da- iini and Hecr, 1867), considered the Dakota kota Group. Based upon studies of recent de- Group of Meek and Hayden (1856, 1861) to positional facies and comparing them to the have been deposited in fresh water. This deter- Dakota, Lesquereux stated: "They are beach mination was based primarily upon the included formations, like those in progress at the present leaf flora. However, Hayden (1867) stated that time along the shore of the North Sea, in Hol- together with F.B. Meek, he had colleceted well- land and Belgium, where the widelv extended preserved marine invertebrate remains mingled mudd\' shores are formed of a soft substance with the leaves of the Dakota Group. Hayden of the red color." Lescjuereux further mention- concluded that the Dakota Group was marine ed that the presenth' forming North Sea beach- in origin. This conclusion was adhered to by es are characteristic and similar to Dakota beds Lescjuereux (1874, and others) and by subse- by being composed of sands borne by the sea quent workers. intermixed with muds borne by the sea inter-

^Departjiient of Botany and Range .Stienie, Brighani Vowig Unl^eI^i^y Bhicham Young University Science Bulletin

Dakota mixed with muds borne by nearby rivers. In responsible for the deposition of the magnitude and mentioning that the leases of some species often Sandstone was of considerable large area. Second, this geo- exhibit a cUimped rather than a random distri- had effect over a the extensive marine advance bution in Dakota strata, Lesquereux (I.e.) postu- logical event was Dakota times. Third, the Dakota Sand- lated: during stone Formation indicates a set depositional of this kind can result only from A distribution characteristics rather than conforming to a dis- the proximity of the trees from which the tinct lithological or paleontological division. leaves liave been derived, and confirms the opinion that the formation of the Dakota From the foregoing discussion, a working whose sur- group is the result of muddy flats definition of the Dakota Sandstone Formation face, raised perhaps in hillocks above water- mav be given. This formation is a Cretaceous limits, and already solid j^round, was cut like sequence of marine, fresh or an immense swamp, here and there inter- time transgressive spersed by rare groups of trees and bushes. brackish water, clastic sediments of various colors (with red, \ellow, and white being com- Lesquereuxs hypothesis, it In support of mon) deposited under fluvial and paludal con- mentioned that within the Dakota should be ditions, often with interbedded lignite, shale Formation there are deposits indica- Sandstone and ash secjuences. This formation can not be brackish water. Evidence for tive of fresh and defined on the basis of paleontological similari- this comes from hgnite seams, fossil assem- ties, although included fossil plants may be cor- blages, stratigraphy and lithology. These sedi- related to some extent. Strata comprising the deposited in embay- ments could have been Dakota Sandstone Formation are related estuaries, and on flood plains (Chaney, ments, throughout their geographical extent in that Repenning and Page, 1956.). However, be- 1954; thev were similarlv deposited as the result of of frequent lignite beds and carbona- cause the invading Cretaceous sea. The Dakota Sand- shales, and the great number of leaf im- ceous stone Fonnation exhibits continuity throughout pressions, which, due to their excellent state of much of the western and midwestern United preservation, do not appear to have been trans- States, authough sediments of this formation ported for any great distance, these deposits from different regions need not be of the same appear to have been laid down in paludal en- age due to the deposition of Dakota strata at vironments close to the sea, similar to those the edge of a coastline which changed with described bv Lesquereux (1874). time. In discussing the nomenclature used in con- Tester (1931) pointed out that a wide variety nection with the Dakota Sandstone (as used bv of names had been used for discussing Dakota Meek and Hayden, 1856, 1861), Tester (1931) strata including Dakota Group (originally used preferred the useage of the term Dakota Stage. by Meek and Harden in 1861 in referring to His reasoning for the application of this term their Formation No. 1 of 1856), Dakota Series, sheds some light on the nature of the beds re- Dakota Stage, Dakota Formation, Dakota Sand- ferred to the Dakota Sandstone. Tester stated stone, and Dakota without an\' additional term. in part: Tester (1931) preferred the use of Dakota Stage which he used svnonvmously with the term Geologic events which ;ire of considerable magnitude, and which have some effect over group, since he restricted the usage of group a large area, or which constitute a normal for the rocks of an entire era. progression of rocks, are considered responsi- The term group or stage usage is useful in ble for deposition rocks comprising the of a delimiting the rocks of more than one closely stage. The widespread marine advance, with related formation when the distinction between its shoreline variations due to minor retreats and advances of the waters and to the lands them is either difficult or unnecessar\'. How- being built out into the oceans, or the migra- ever, later workers (Jcnney, 1899; Darton, 1905; tion of faunas and shifting of ocean currents, Stanton, 1905) showed that the Dakota Group all have tlieir effect on the character of the of the Black Hills and Rock\- Mountain regions rocks. The rocks deposited under such conditions on an extensive scale, as they were dur- could be divided into Lower and Upper Cre- ing Dakota time, are classed as a stage. It taceous formations; these workers restricted the miglit be said tliat a stage is indicative of a Dakota Sandstone to formational status. Fur- .set of conditions of rock deposition rather than thermore, the original Dakota Group from the of a distinct lithological or paleontological division. midwestern United States is similar strati- graphicalK' and floristicalh' (although some From Tester's discussion, several important variation mav be noted in floras from different concepts arc apparent. First, the geologic event regions) to the restricted Dakota Sandstone Biological Series, Vol. 14, No. .3 D.\kut.'\ Sandstone Flora

UTAH

ARIZONA

Fic. 1. Index map of collecting localities. HiuciiAM VoLNC University Science Bulletin

Formation of the Black Hills and Rock\' Monn- the Burro Camon Formation, and a somewhat from the tain regions. In light of this, the term Dakota at\pical Dakota flora was described Sandstone Formation will be used by the author Dakota Sandstone Formation. This Dakota flora of ferns with for discussion of Dakota strata. is at\pical in that the incidence angiosperms is high. Geolcx;y. -Stanton (1905) in working with the Jurassic and Cretaceous stratigraphy near Jurassic Formation and its relationship with the Westwater, Grand County, Utah, is similar to Comanche Series and the Dakota Formation in that of Montrose Countv, Colorado. The Jur- Southern Colorado, New Mexico, and Okla- assic Morrison Formation from this region of homa, demonstrated that the Dakota Sandstone Utah is overlain b\ the Lower Cretaceous of this region, as originally defined, contains Cedar Mountain Fonnation. This formation is both Lower and Upper Cretaceous strata. Thus, conelated to the east with the Buno Canyon Stanton pointed out that the true Dakota Sand- Formation. However, no plants have been col- stone Fonnation of these widely separated re- lected from the Cedar Mountain Formation of gions overlies Lower Cretaceous deposits which Grand County. This Lower Cretaceous for- overlie the Morrison Formation. mation is overlain by the Dakota Sandstone Similar conditions have been observed in Formation (Plate 3.) This formation from this several other regions of the western United region is composed of three lithological units. States. In Montrose County, Colorado, the Da- The basal unit is a massive, buff-colored sand- kota Sandstone Formation occupies a position stone of approximately thirty feet in thickness. between the Jurassic Morrison Formation (Mc- This unit is overlain bv a shale-coal-sand se- Elmo Formation of Coffin, 1921) and Upper quence which is approximately thirt\' feet thick. Cretaceous Mancos Shale Formation. Careful Fossil plants collected near Westwater are ob- examination of these strata (Coffin, 1921) in- tained from this unit. This shale-coal-sand unit dicated that the Jurassic Morrison Fonnation is in turn overlain by a massive buff-colored is overlain 1)\ a Lower Cretaceous sequence. sandstone unit. This is the uppermost unit of This sequence was given formational status bv this formation near Westwater, and it is ap- Coffin (1921) who named it the Post-McElmo proximately thirty to forty feet in thickness. Formation. Stokes (1948) changed the name of This formation is overlain b\- the Upper Cre- this Lower Cretaceous setjuence to the Burro taceous Mances Shale Formation. Most leaf Canyon Formation. compressions collected from the Dakota of this Plants of this Lower Cretaceous setiuence in approxi- Colorado were studied by Brown (1950). Brown region are obtained from an ash seam concluded that floristic evidence agreed with mately forty to forty-five feet beneath the Grtj- the findings of Coffin (1921). A Lower Cre- phiiea ncubernji zone in the overlving Mancos taceous flora was described bv Brown from Shale.

THE AGE OF THE DAKOTA SANDSTONE

Much of the earliest work concerning the ing workers, and the only subsequent contro- Dakota Sandstone Fonnation was concerned ex- versy has been concerning the proper Creta- tensively with the age of these rocks. Briefly, ceous epoch to which Dakota strata should be the Dakota was the first considered as a Cre- assigned. taceous group (Meek and Hayden, 1S.56, 1858). An accurate ascertainment of the age of the This interpretation was refuted b)- Hawn (1858) Dakota Sandstone Fonnation (whether early or who supposed that Dakota strata belonged in late Cretaceous) has been complicated by two the Triassic. Heer (1859, 1861) proposed that |)r<)l)lems. The first concerns a poor usage of the Dakota was of Tertiar\ age, and Marcou the term. "Dakota flora." This term has been (1858) stated that the original Dakota was com- used loosely bv some workers, and as pointed posed of rocks of both Tertiarx' and Jurassic out by Berry (1920), "Any Cretaceous formation age. Later work (Marcou, 1864; Cajiellini and containing dicot\ledonous leaves and known or ." Heer, 1867) demonstrated that Dakota rocks thought to be older than the Benton . . was indeed belong to the (Cretaceous, as originalh' said to contain a Dakota flora. In other words, postulated bv Meek and Hayden. This age de- Dakota flora became an adjectival term rather termination has been adhered to b\- ail follow- tliaii dcliiniting a flora which had been ob- BioLooicAi. Series, \'()l. 14, No. 3 Dakota Sandstone Flora

within the formation or from evolutionary changes in plant communities over a period of time svnchronou.s with deposition. An alternative to the proposal of Berry m (1920) is that in delimiting a true Dakota flora, »' IT only plants from the Dakota Sandstone Forma-

tion be considered. That is, the onlv valid application of the term flora -.iJ9. Dakota can be in the .,^ discussion of plants from the Dakota Sandstone sensu stricto. It is neither valid nor prudent to /--• delimit a Dakota flora, merclv in the sense of a Cretaceous dicotyledonous flora, from anv ^*^-, other geological entity. '.r^ This ._. -w suggestion, however, leads to a discussion of the second problem which is some-

what less easily treated. It is that the Dakota Sandstone is a time transgrcssive formation and ..:^

V>.>.^~,*1

Fig. 2. .K. Southeast view down canyon towards West- water, Grand County, Utah. The Dakota Sandstone Formation forms the hills and slopes in the fore-

ground and is exposed along the roadcut. Original »i^fb»n collections from the Dakota Sandstone of this region

were made in the roadcut ( A ) . The Mancos Shale Formation o\erlies the Dakota in this region. B. Close-up of collecting site (A) in Fig. 1. This site \ielded the best colections of fossil plants from this area.

tained from the Dakota Sandstone. This problem became so acute that Berrv (1920) further stated, "It has become increasingh' clear of late years that Dakota flora was not a unit and had no precise stratigraphic value." Berrv (1920) proposed that in order to eliminate this problem, a true Dakota flora mav be defined as "mean- ing thereb\ the equivalent of that of the Wood- bine Formation of Texas, and those of corre- ." sponding age elsewhere. . . This interpretation, houever, has one inherent difficulty. That Fk;. .3. A. Overview of Dakota Sandstone Formation is, it does not allow for the possibility that showing massive upper sandstone unit resting upon within the same formation, fossil assemblages sliale-coal-sand imit. The arrow points to a collect- ma\ differ sufficienth' to render their com- ing excavation. B. Close-up of excavation in Fig.

1 illustrating lithology of shale-coal-sand unit. The |iarison with the flora from another formation upper coal (B) and unfossiliferous ashes (A and C) difficult. This ma\' result either from a non- have been removed in this excavation exposing the random geographical distribution of fossils fossiliferous ash upon which the worker is kneeling. .

6 Bni(;nAM VouNc; University' Science Bulletin was deposited over a range of time that wit- T'^<^arJ-- nessed a vast floristic change in the dominant --^9 vegetation of the earth. That is, although all formations are time transgressive, many are deposited within a period of time wherein little change in the contemporaneous flora or fauna occurred, and therefore the included fossils of that formation would be expected to exhibit homogenitv. However, the Dakota Sandstone Formation was deposited over a period of time when the predominant vegetational type of the earth changed from a fcrn-g\'mnosperm alliance typical of the older Mesozoic to an angiospemi dominated flora typical of the Late Cretaceous,

Tertiary, and present times. Therefore it is possible to collect within this formation, fossil assemblages which appear to be indicative of "7 different ages. This is particularly the case when the Dakota Sandstone Formation from I west and east of the Rockv Mountains are compared, and the floras from the western portion of this formaton appear to be older than their counterparts from the east. Further discussion on both microfloral and megafloral evidence bearing on this situation will be considered later. Earliest detailed estimates (Lesquereux, 1874, 1883, 1892) concerning the age of the

Dakota Sandstone Formation placed it as Ceno- manian. This concept has been generally adhered to since that time, although some geol- KiG. 4. A. Roadcut exposure of Dakota Sandstone gists have disagreed with it based chiefly upon Formation. The upper massive sandstone unit (A) stratigraphy. In connection with this, Twen- of this formation from this region rests upon the hofel (1920) assigned the Dakota to a Lower middle shale-eoal-sand unit ( B ) . C dehmits an Cretaceous age. Tester (1931) in studying the unfossiliferous ash, and D represents the chief fossiliferous B. Ix'tween Dakota type locality of the Dakota Sandstone consid- ash seam. Contact Sandstone and Cedar Mountain-Morrison Formation. ered it to be older than Ccnomanian and placed The Dakota forms tlie slope and ledge ( A ) which Washita-Kiowa-Mentor-Dakota rocks of Kansas rests upon the slope of tlie Cedar Mountain-Mor- at the base of the Cretaceous, with the Da- rison Formation ( B ) kota being at least as old as the Mentor Forma- tion of this region (midAlbian). However, the that the Dakota was of Upper Cretaceous (Ceno- conclusions of Twenhofel and Tester were manian) age and could be correlated with the drawn based upon stratigraphic evidence alone Woodbine Formation of Texas. However, Berr\' and are not substantiated by floristic evidence. (1922) later considered the Woodbine Forma- Cobban and Heeside (1952) assigned the tion to be of Turonian age, although still to be Dakota (Jroup to various ages ranging from correlated with the Dakota Sandstone Forma- early Aptian to middle Cenomanian. However, tion. This determination was based upon a de- the lower members of this group (Lakota Sand- tailed anahsis of the flora of the Woodbine stone and Fuson Shale of Darton, 1905) were i'ormation from Lamar Count\', Texas, where-

assigned to ages earlier than late Albian, and in it was noted bv Berry that this flora had the restricted Dakota Sandstone Formation was three species in common with Turonian floras considered by these workers to range from late of Europe. Albian through middle Cenomanian. MacNeal (1956) restudied the Woodbine Berry (1920) was opposed to the detenni- flora from collections made in Denton County, nation of Twenhofel (1920) tliat the Dakota Texas. Based upon examination of more and Sandstone Formation was of Lower Cretaceous better material. MacNeal disagreed with the age. In this paper Berry cited floristic evidence age determination of Berry, and a.ssigned the HioLOGiCAL Series, Vol. 14, No. 3 Dakota Sandstone Flora

Woodbine flora to a Cenomanian age. MacNeal Hedlund (1966) also confirmed the Ceno- did, however, agree with Berr\- (1920) that the manian age of the Woodbine Formation. In

Woodbine flora is closeh- related to the flora this paper, Hedlund treated manv species of of the Dakota Sandstone. fern and g\innosperm palvnomorphs indicating A Cenomanian age for the Woodbine was that this segment of the flora, even though also agreed upon b\' Stephenson (1952) based poorlv represented by macrofossils, nonetheless, upon a stud\' of the invertibrate fauna from this represents an important floristic component of formation. the Woodbine. Brown agreed with Berry that (1952) (1920) Pierce (1961), Hall (196.3), and Agasie (1967) the Woodbine flora is verv similar to the flora in reporting on microflora! assemblages from of the Dakota Sandstone of Kansas. One appar- the Dakota Sandstone Formation of Minnesota, ent difference, however, was the conspicuous Iowa, and Arizona, respectively, assigned this absence of g\mnospermous species in the formation to a Cenomanian age. The present Woodbine flora. This difference was elimin- author agrees with this assignment ated by MacNeal (1958) in his Woodbine studies Cenomanian and recenth' b\' Hedlund (1966) in his palv- of the Dakota Sandstone Formation, although nological studies on the Red Branch Member in part this formation mav be of late Albian of the Woodbine. age (Cobban and Reeside, 1952).

A CONCEPT OF A DAKOTA FLORA

It was recognized from earlier plant col- able to the modern angiospermous vegetational lections from the Dakota Sandstone Formation type were exceedinglv common and comprised that plants from this formation were rather by far the dominant part of Dakota vegetation. closely related to plants growing on the earth Indeed, Lesquereux (1874) noted that dicot at the present time. Fossils from these collec- species represented in the Dakota floras could tions were representatives of dicot\'ledonous be referred to "genera to which belong most families and genera most of which are still of the living arborescent plants of this country represented in the earth's flora. As Meek and (North America) and of our present climate." Hayden (1859) commented, Dakota fossils "be- When these facts were considered bv Les- long to a higher and more modern tvpe of quereux (1874), he wrote a section dealing with ." dicotyledonous trees. . . the disconnection of Dakota floras with ante- As more collections were made from this cedent types. In this chapter, Lescjuereux wrote: formation throughout the midwestern United ... it is evident that the flora of the Dakota States from such diverse localities as Kansas, Group is as widely dLsconnected from that of Nebraska, North Dakota, South Dakota, Wyo- the Jurassic, even of the Lower Cretaceous, ming, New Mexico, etc., a floristic picture of or as distinctly original, as are the flora of the Dakota began to evolve. This picture was the Carboniferous compared to that of the Devonian, or the Permian types compared to essentially that of Meek and Hayden (1859), and those of the Cretaceous. later of Lesr^uereux (1874, 1883), that the flora of the Dakota was essentallv a modern tvpe of Later studies (Lestjuereux, 1883, 1892) indi- flora without many fossils representing ante- cated that although ferns and gymnosperms cedent floristic types. As mentioned by Les- were more common than indicated bv earliest quereux (1874) only one specimen collected to studies, these floral types were still present in that time could be referred to the Cycadophyta limited numbers. Thus, Lesquereux (1892) in {PterojiJiijUtim (?) lunjdenii), and even this ref- his final work on the flora of the Dakota Group erence was doubtful. Conifers were rare from summarized the relative importance of the dif- these early studies, both in number of species ferent components of this flora. Of a total of and number of specimens, and represented a 460 species identified from the Dakota Sand- very limited portion of the flora. Likewise, ferns stone flora, Lescjuereux recognized six species were few in number and appeared to make up of ferns, representing 1.3 percent of the total a ver\- limited part of the total Dakota vegeta- flora; twelve cycadean species representing 2.6 tion. percent; fifteen species of conifers represent- On the other hand, even from earliest ing 3.5 percent; and 437 species of angiosperms studies, it was apparent that plant fossils refer- representing 92.6 percent of the total flora. )

8 BmciiAM Voi'Nc: Univkhsitv Science Bulletin

In the liglit ot the above discussion, it may work by Cockerell (1916) and Berr%- (1919a) in- be summarized that the prevailing early con- dicate the presence of matoniaceous ferns from cept of the flora of the Dakota Sandstone For- this region. Ferns referal^le to this famih' are

mation was that it was essentially a modern representative of an earlier flora! t\pe, as they flora composed principalK of dicotyledonous are common in the European Wealden and leaves related to modern arborescent genera. other Middle Mesozoic strata. Matoniaceous The apparent paucity of antecedent floral t\pes ferns are uncommon from rocks younger than (Jurassic-VVealden ferns and gyinnosperms Earh' Cenomanian, and are unknown from

common in earlier Mesozoic floras is note- Tertiarx' strata. Brown (1950) made further worth\', although somewhat enigmatic to some studies on tlu' plants from several localities in earh' paleobotanists. It must be remembered, the Dakota of Montrose County. This author however, that this concept grew out of studies also treated matoniaceous ferns, as well as ferns by Lesquereux and others of macrofossils from of the famih' Gleicheniaceae, which are also the midwestern portion of North America and indicative of a preangiospermous floral type. was drawn without the aid of data from two Several other ferns from the Dakota of this sources to be added later. These two sources region were also chscussed b\' Brown. The are microfloral anahsis of the Dakota, and author has had occasion to study the Dakota megafloral analysis of this formation from many collections from Montrose County reported by new localities, especially those further west Brown. It is apparent that the fern families than original collection sites. Matoniaceae and Gleicheniaceae were very well As more data accumulate concerning the developed in this region during Dakota times. Dakota Sandstone Fonnation, particularly data Indeed, ferns of these two families are so com- dealing with regions west of the Rocky Moun- mon that they make up the predominant part

tains, it becomes clear that a valid concept of the fossil vegetation, although they are col- of a Dakota flora must be modified some- lected in association with angiospemious forms what from early views. While this earlv con- which, as mentioned bv Brown (1950), cept is essentialh' for megafloral assemblages . . . include chiefly species that occur in the from that portion of this fonnation east of the large flora described by Les<]uereu.\, New- berry and others from the brownish Rocky Mountains, it does not fit well for the Dakota •Sandstone of Kansas and Nebraska, in the formation as a whole. In this respect, it is sig- Woodliine Formation of Texas, the Dakota of nificant that the incidence of species repre- the Black Hills, South Dakota, and the Upper sentative of the antecedent Jurassic-Wealden Cretaceous rocks of Greenland. vegetational type, particularh' Middle Meso- Therefore, during Dakota times in southwest- zoic ferns, is much higher in megafloral assem- ern Colorado, the vegetation was composed of blages from several localities in the Dakota typical Dakota angiospemious species, but with Sandstone Formation west of the Rocky Moun- a large proportion of plants representative of tains. That is not to say that dicotyledonous a more ancient floristic type. species are not present at these localities. The\- A somewhat similar situation existed in are not only present, but they are directly re- Grand County, Utah, during Dakota times. Sim- lated to the dicot\ledonous species from this ilar to the fossils from Colaroda, the Dakota formation further to the east. Indeed, at some flora from this part of Utah is comparable to of these collection sites, angiospemious fossils Dakota floras from other localities. However, comprise the dominant floristic component of it is both significant and apparent that the the fossil assemblage. H()we\er, from man\' flora from the Grand Count\ Dakota Sand- localities in the Dakota Sandstone Formation stone Formation is ver\' much dominated b\' in Utah, Colorado, and Arizona, fossils repre- ferns representative of the older Jurassic- sentative of the older antecedent vegetational W'ealden vegetational t\pe. Indeed, it is al- type are relativel\- more important than their most impossible at some localities in Grand counterparts east of the Rockies, both in num- C'ounty to crack open a rock without exposing ber of species and number of specimens pre- at least one specimen of a fern representative sent. Indeed, at some western localtities, repre- of the families Matoniaceae or Gleicheniaceae. sentatives of this ancient floral l\pe dominate Dicf)tyledonous species are reduced both in the floral assemblage. number of species and number of specimens The Dakota flora from Montrose County, present. Gxmnosperms represent a rather minor Colorado, represents an admixtur(> of repre- part ot the flora from this region as far as sentati\'es of the Jurassic-Wealden vegetational number of species present, although quanti- type with modern angios|)ermous forms. Farl\- tatively they are represented b\- large amoimts .

Biological Series, Vol. 14, No. 3 D.\kot.\ S.\Nn.sTONE Floh.\ of .silicified wood. Therefore, the Grand Count) from microfloral anahsis of two localities in

Dakota flora mav be characterized a.s a fern- the Dakota from Minnesota and Iowa respec- dominated, fern-angiosperin-g\inno,sperm alh- tivelv, that ferns and gsmnosperms represented ance, .similar to what could be expected during more important components of the eastern Da- a time which directly preceded the much dis- kota flora than indicated by the megafossil re- cussed "population explosion" of angiosperms cord. With respect to this. Pierce reported 24 during Cenomanian times ( Seward, 1927 ) species of pahnomorphs representative of an- This same vegetational aspect has been ob- giospermous species, 36 species of gvmnosperm- served b\' the author from several other Da- ous pahnomorphs, and approximately 20 kota localities of the western United States. species of fern pahnomorphs. Based on this, Floras from Longhouse \'allev and Kayenta, Pierce postulated that angiospermous species Na\ajo Countv, Arizona, both exhibit character- are not as important in this formation as indi- istics discussed above in connection with Colo- cated bv the megafossil record. However, a rado and Utah Dakota floras. That is, ferns re- more precise conclusion would be that rather presentative of the old Jurassic-Wealden vege- than the angiosperms representing a less im- tational t\pe dominated the Dakota vegetation ])ortant floral component, the gvmnosperm-fern of these regions, and angiosperms represented floral component represents a more important a rather limited vegetational type. part of the flora than previously thought. The Dakota Sandstone Formation from Coal From the above discussion, a more accurate Can\on, Coconio Countv, Arizona, as discussed concept of the flora of the Dakota Sandstone bv Agasie (1967) exhibits similar floristic char- Formation as a unit may be formed. In western acteristics. Microfloral analysis from this region America, the Dakota flora is a fern-angiosperm indicates that Dakota vegetation was composed alliance with a relativeh' small gvmnospermous of a fern-angiosperm alliance with a minor component. At several localties west of the g\ mnospermous component. Fern spores repre- Rockies, ferns dominate the megaflora as well sent the dominant microfloral component, es- as the microflora. To the east, this flora pecialh' fomis related to the famlv Schizaceae. changes character somewhat, becoming an an-

This fern famil\- is often represented in pre- giospenn-dominated flora with a fern and gym- angiospermous floras (Harris, 1961), and pre- nosperm component. This fern-gymnosperm ]iminar\' collection of megafossils from the area component was originalh' thought to be of very also indicates the presence of manv ferns re- minor importance in the eastern Dakota, but lated to an old Jurassic-Wealden vegetational recent microfloral studies (Pierce, 1961) have tvpe. demonstrated that this component of Dakota

In addition to the data presented above, it vegetation was more important than had been has been shown by Pierce (1961) and Hall (1963) previousK' supposed.

PALEOECOLOGY

The plants from the Dakota Sandstone For- of the same species, and laterally the species mation near Westwater are preserved in a light change abruptlv within a distance of a very tan ash la\er from five to ten inches in thick- few feet. In certain outcroppings of the ash, ness. This laver was deposited directly upon a dense mats of leaves are found within a rela-

coal seam and is overlain b\' another. These tiveh' small area. These mats are normally plants are extremely well preserved with manv found in the lower portion of the ash, and if of the leaf compressions illustrating cuticle, one traces them lateralh', the\' appear to follow vascular tissue, and reproductive structures. In definite channels indicating possible stream addition, manv of these fossils are disposed in channels. the strata in such a manner as to indicate that Peels of the foliage of the channel areas the plants were preserved in growth position, were prepared to aid in identif\ing the angio- suggesting that deposition of the ash was rapid. sperms. On one of these peels a well-preserved

The ash la\er is uniform in color and com- diatom and several spores of either algal or position throughout, with the exception of a fungal origin were found, indicating the likeli- one- to two-inch portion directK- beneath the hood that the depositional environment of upper coal seam. Lateralh throughout the ash, matted leaves in these areas was aijuatic.

distribution of the plants is differentential. More detailed study of the fossihferous ash Thus, within a local area, most of the plants are brought to light several other important factors. .

BmcnAM VoiNx; Univeksity Science Bulletin 10

plant- In all areas lu-ar Westwalcr wIhtc the beariiii; asli lias hi-en exainiiu'd, tht" contact between the ash and the nnclerl\ ing coal seam is very sharp and easily distinguished (Fig. 5). In addition, the lowemiost portion of the ash contains large nnmbers of root and rhizoid- like structures, often in such profusion as to form dense mats of these structures (Fig. 6). Leaves are scarce in the lower portion of the ash with the exception of poorly preserved, often broken specimens. One of the more important identifiable fossils common in this region of approximately one inch in thickness is the rhizome of Equmtum lijeUi Mantel with attached tubers. Immediatelv above this rhizome region is a horizon containing numerous well-preserved leaf impressions. This region contains foliage belonging almost exclusivelv to the three fern genera Matotiklitiin, Astnilo])tcris, and Gleich- enia. This zone is approximately five to eight inches thick and is literally full of beautifully preserved specimens. At the upper portion of this dense foliage region, the color and composition of the ash changes somewhat (Fig. 5). The ash becomes noticeabh darker in color and somewhat silty- carbonaceous in composition. Many of the fos-

sils found in this region differ taxonomically from those found lower in the ash. While the

three genera Matonklitim, Astraloi)t('ri.'i and Fig. 5. A. Collectins; .site e.xhibitiiig characteri.stic lith- Gleichenia are still present as leaf impressions ology of Dakota Sandstone near We.stwater. The sandstone unit forms the chff, and in this region, thev are not as dominant as they upper massive tlie weathered shale-eoal-sand secjuence forms the were in the lower portions of the ash. tahis slope in the foreground. B. Close-up of fossils in region represent Plant this upper fossiliferous a.sh seam illustrating rhizome region

taxonomic , di\'ersit\' region ( ) several different genera. The greatest ( 1 ) , leaf region ( 2 ) and 3 diversity in the Westwater flora occurs within this zone. Significantly, Equisetuin rhizomes structures, rhizomes, partially decomposed reoccur in this upper region after being noticc- leaves, stream channels, etc., occurs. Above this

ablv rare in collections from the middle portion is a region extremely rich in matoniaceous, of the ash. gleicheniaceous, and astralopteroid fossils which The contact between tlu' ash and the upper appear to be in growth position. This evidence coal seam is not as sharp as the lower contact would tend to support the hypothesis that the (Fig. 5). From the point where the ash be- ash was deposited rapidly upon a coal swamp comes carbonaceous in nature and diversitv in which was composed chiefly of ferns of the the flora appears, the ash laver graduallv be- three genera Matonidium, Gleichenia and Astni- comes more carbonaceous until within a dis- lopteris. If this indeed occurred, the rhizome tance of one to two inches it grades into a well- region would contain mostly rhizomes and roots defined lignitic coal. of these three genera. Several explanations for the above described The upper portion of tlie ash where the phenomena have been considered and rejected. diversitv appears contains plants which may rep- The sharp contact between the lower coal and resent forms that began to grow upon the

ash indicates that the ash fell directlv upon previously deposited ash. That is, the plants of a coal swamp and preserved the jilants that the upper part of the ash represent forms that were growing at the time. Further evidence in repopulated the swamp following ash deposi-

supjiort of this is noted when it is recalled that tion. Tlie likelv reason for the diversity in taxon-

at the base of the ash a region rich in rootlike omy of these plants is that they are forms BioLOGicAi. Skbies, Vol. 14, No. 3 Dakoiw S.^ndsionk Floh.^ n

REGION OF DIVERSITY

RHIZOME SAND REGION

Fk.. 0. I.^ Representative forms foiiiul in rt-f^ioii of diversity: Conioptcris (a); Asplcniutn (b); Equisctum (c); and Ctmlophlchts (d). 2. Single specimen collected from leaf region illustrating Gfetc/icnifl and A/rttomV/iHni. 3. Rhizome and rootlike structures collected from rhizome region. 12 Bhiciiam Vouno University Science Bulletin from serai communities in secondary succes- regions shows no difference when examined sion. It is Iikel\- that the climax vegetation microscopicalK-. However, the ash of the region growing in the coal swamp was a Matonidium- of diversity is much more silty and carbo- Glcicheniu-Afitraloptchs association. Further evi- naceous than the ash of the two lower regions. dence in support of this conclusion was obtain- This evidence would tend to support the theory ed b\' the author from stud\' of mascerations of that the jilants of the rhizome and leaf regions the lower and upper coals. Even though paly- were pri'siTved in one ash tall. The silty carbo- nomorphs from these coals were poorly pre- naceous region (region of diversity) represents served, it was evident that matoniaceous fern a region of increased silt and carbon deposition spores were the dominant microfossils from due to the initial deposition of enough soil to both, indicating that species of Matonklhim support plant growth, population of this soil were prevalent both prior to and subse(|iient to b\' new plants (secondar\' succession), and sub- ash deposition. sequent carbon deposition as these plants died To test the above hypothesis, thin sections and were deposited. These plants probably rep- of the ash were made from the three discussed resent stages in serai succession toward the regions (rhizome region, leaf region, and diver- climax coal swamp vegetation of Matonidium, sity region). The ash of the rhizome and leaf Gleiclienia and Astmloptcris.

PALEOCLIMATE

It is sometimes possible, particularly when margins, 44 percent are over 10 cm long, and working with late Mesozoic and Tertiary paleo- 74 percent are thick in texture. From these floras, to reconstruct the presumed climate un- data. Chancy concluded that the Dakota flora der which these fossil floras existed. Two basic lived under conditions "more subtropical than metliods used in determination of paleoclimates temperate." Chancy further pointed out that are the comparison of fossil genera to their 72 percent of Dakota leaves exhibit campto- nearest living relatives existing under known drome venation patterns which, according to climatic conditions (extrapolation through uni- Chancy, also indicates that the climate during formitarianisin), and the detennination of dicot- Dakota times was tropical or subtropical in yledonous leaf characteristics and comparison nature. (>hanev suggested that this angiosperm of these characteristics with those found in flora existed in a lowland forest, and that some species growing in known climatic conditions. highland regions were to be found nearb\'. It was first recognized b\' Bailey and Sin- Pierce (1961) discussed the Cretaceous cli- nott (1915) that leaves of arborescent dicot\le- mate of Minnesota based upon a fossil pol- donous species growing in tropical regions ex- len assemblage from this region. He concluded hibit a higher incidence of entire margins tlian that the flora could be characterized as a coni- do their counterparts growing under non- ferous rain forest growing under warm tem- tropical conditions. These authors pointed out perate climatic conditions similar to conditions that there is a correlation between the number found ]iresently in the Pacific Northwest of the of entire margins of arborescent dicot species United States or the coastal regions of eastern to climatic conditions under which these species Asia. This was based upon an unusualK- high exist. Thus, from sevent\' to ninetx' percent of tre([uenc\' of g\mnospermous pollen encounter- tropical wood\' dicot species exhibit entire leaf ed which. Pierce argued, could not have come margins, whereas only twenty to forty percent from nearby upland regions. Based upon this of wood\- dicots of cold temperature regions ex- information. Pierce agreed with Les(iuereux hibit entire margins. Further studies (Sinnott (1S74) who suggested that the Dakota flora was and Baile\-, 1915; Chane\- and Sanborn, 19.33) similar to the modern flora growing between pointed out that large leaves, thick leaves, com- 30° and 45° north latitude. However, Lesquer- pound leaves, and leaves with attenuated apices cux never assumed that the Dakota flora was are more abundant in tropical regions, decreas- a coniferous forest, or that it even contained ing proportionately into temperate regions. a large number of g\mnosperms. Indeed, just Chane\- (1954) applied these characteristics the opposite was the case as Lesquereux (1874) in analw.ing 200 species of leaves from the discussed the noteworthy absence of g\mno- Dakota flora in an effort to determine the paleo- spermous and fern species from the Dakota climate of Dakota times. He noted that 73 per- flora. The work of Pierce, however, does point cent of the leaves from this flora exhibit entire out that both the g\ mnospennous and the fern )

Biological Sehies, XOi.. 14, No. .3 Dakota SANDsroNK Flora 13 component of the Dakota flora are larger than sented in the Grand Cx)unt\- flora. This family indicated h\- nicgafloral anahsis. is monotypic containing the genus Di])teris, Hedlund (1966), in considering the pollen which is presently restricted to fiye species all flora from the Red Branch Member of the of which are found in the Indio-Malayan re- Woodbine Formation which is closeh' related gion of the world (Bower, 1923). Thus, this to the flora from the Dakota Sandstone, com- famih' also may be cited as an excellent ex- pared component members with their modern ample of a tropical or subtropical climatic in- counterparts and concluded that the climate of dicator. this region during \\'oodbine times was warm In addition to the aboye evidence, most ob- temperate to tropical. His determination was served angiospermous fossils exhibit entire leaf based upon the high incidence of palynomorphs margins and two species show long attenuated related to modern plants which are presently apices (drip points) which are thought to func- distributed chiefly in wami temperate-sub- tion in facilitating the run off of excess mois- tropical regions. ture. As mentioned previously, these character- A Dakota pollen flora from Coal Canvon, istics are found more frequently in tropical re- Arizona, indicates subtropical to tropical cli- gions, and thus, the Dakota angiosperms from matic conditions for this region during Dakota Westwater substantiate the conclusion that the times (Agasie, 1967). This is based upon rela- Westwater Dakota flora grew under subtrop- tivel\' high frequenc\' of pahnomorphs from ical to tropical conditions. genera related to extant taxa growing under Topography at the time of deposition of the wet tropical or subtropical conditions. The rela- shalc-coal-sand unit of the Dakota Sandstone tiveh' high frequenc\' of g\mnospermous pollen Formation of this region was likely a broad present in this flora indicates that well-drained, swampy mudflat near the shore of the Ceno- low upland regions existed near the lowland manian Sea. Pockets of dense vegetation com- swamp\' depositional basins. posed basically of ferns and Equisetum de- Several species and hundreds of specimens veloped in some of these regions, often accumu- of Matonidium are present in the Westwater lating enough volume, under proper conditions, flora. These ferns are probablv the best cli- to comprise a large component of the coal de- matic indicators present in this flora. These fos- posits which occurred between sandy or shale sils are related to the fern famih- Matoniaceae sequences. Well-drained highland areas existed which is presently a monot\pic (Bower, 1923) nearby with gvmnosperms and angiosperms or ditypic (Holtman, 1947) family containing present. Streams ran from the upland regions the genus Matonio. This genus is extremely into the lowland swampy areas carrying leaves limited geographically and climatically in the and other debris in various stages of decom- present world's flora and grows only in the position. The vegetative deposits from these humid tropical upland regions of the Malayan streams (gymnosperm and angiosperm leaves,

Peninsula. As mentioned by Mahabale ( 19.54 and likely wood) probably formed a large por- matoniaceous ferns are among a select group of tion of the coals, with the remainder being ferns that are accurate climatic indicators. added from the ferns and associated plants of Gleicheniaceous ferns are well represented in the fern swamps. the Westwater flora, although these fossils are Following a period of time when deposi- somewhat less yaluable than matoniaceous ferns tion occurred as outlined above, the shoreline as climatic indicators. Of 80 extant species the of the sea changed, and the shale-coal-sand se- majority are of tropical distribution, although quence in the Dakota of the Westwater region some species extend well into southerK- lati- was overlan bv a massive sand sequence. Fol- tudes (Bower, 1923). lowing this, the entire formation was overlain The fern famih' Dipteridaceae is also repre- by deposits from the Mancos Sea.

SYSTEMATICS

Division ARTHROPHYTA to use Equisetum since "No morphological difference has ever been proved between Equi- Genus EQUISETUM L. ." setum ami Equiselites. . . This statement is Harris (1961) discussed the usage of Equi- completely accurate, and furthermore, the fos- setum and Equisetitcs for fossils exhibiting sim- sil and living forms exhibit striking morpho- ilarities to the extant Equisetum. Harris elected logical similarit). In view of this, the author Bhic;h.'vm Vouno University Science Bulletin 14

highly branched, although a rhi- has tollowed Harris (J961) and others in using species was attributed to this species Equisetiim for placement of fossils wliich ex- zome (juestionably 179, Fig. is very much similar hibit characteristics similar to those of the ex- (op. cit., PI. 8) collected from Westwater. tant genus. to rhizomes Bell (1956) illustrated tuber-bearing rhizomes of Equisetites hjelli (Mantel) Unger Equisetum hjelli Mantel (Equisetum hjelli Mantel). These rhizomes dif- Figs. 7-3, 7-5; 15-5. 15-9. fer from the Westwater specimens by being branched and exhibiting strongh' ribbed inter- 1889 Equisetum manjhmdicum Fontainc-U.S. nodes. Other differences are the whorled dis- Geol. Surv. Mon. 15, p". 65, Pi. 170, Fig. 8. position of tubers and the apparent lack of 1893 Equisetum hjeUi Mantel, Dawson-Trans. bracts subtending tubers on the Canadian Roy. Soc. Can. v. 10, pt. 4, p. 83, text-Fig. 1. specimens. Howe\er, the dimensions of the Canadian material compare \sell with those 1956 Equisetites lijeUi (Mantel) Ungcr, Bcll- noted for Westwater specimens, although the Geol. Surv. Can. Mem. 285, p. 76. PI. 28, Figs. tubers are commonh' somewhat larger in the 5-8. latter. unbranched, DESCRiPTiON.-Rhizome horizontal, Other citations of E. hjelli are incomparable to slighth ribbed, 2-4 mm in diameter, smooth to the Utah specimens since onl\' aerial stems branches at inter- bearing short tuber-bearing are described. Thus, Schenk (1871) described tubers ovoid to globose, vals as close as 8 mm; aerial stems of £. hjelli as being much branch- subtended b\- leaflike commonh- wrinkled, ed, and with larger dimensions than would be wide, borne in bracts, 13 mm long by 6 mm expected for aerial stems of the Westwater pairs; upright stems unknown. Equisetum, and Fontaine (1889) noted that stems of E. hjelli from Occurrence.—Dakota Sandstone Formation near characteristics of aerial agreed closely with those of this species Westwater, Grand Count)-, Utah. Virginia from Europe as noted by Schenk (1871). Repository. -Brigham Young University, BYU 1807-1810.

Discussion.—Many specimens of Equisetum Division FILICOPHYTA have been collected from the Westwater flora. Genus ASPLENIUM L. These specimens may be easily identified by although positively identi- their attached tubers, This genus is rather large and is well repre- is absent. Specimens obtain- fied stem material sented both in the modern and fossil flora. for most ed from this flora are collected the Bower (1923) reported Asplenium to contain por- part from the lowermost and uppermost 429 extant species, and Jongmans (1957) listed tions of the ash. 96 fossil species of this genus with three vari-

It is rather difficult to assign this plant to eties and listed ten as Asplenium sp. In addi- anv known species since no aerial stems have tion, Asiilenites has been used for placement been observed. However, rhizomes of Equise- of asplenioid ferns resembling the modem As- tum are not uncommon in the fossil record, and plenittm; this genus contains several additional the Westwater material rather closely resembles species. some described specimens. Most of the fossil species of Asplenium are Fontaine (1889) described Equisetum vir- based upon sterile foliage, and undoubtedly ginicum based upon both aerial stem and rhi- man\ arc s\nonvrns. This problem is further zome material. The aerial stem was highly complicated bv the fact that sterile foliage of branched, and tubers (called imperfect buds by this genus rather closeh resembles foliage of Fontaine) were borne individually along the several other fern genera such as Anemia, Omj- rhizome. This specific name was used again chiopsis. Sphenojiteris. AcrostirliO])leris. Coni- later b\- Fontaine (1899) for placement of a opteris, and Dicksonia. There is no adeciuate specimen from the Black Hills Lower Cre- wa\' of determining between some species of taceous. Bell (1956) noted that in overall mor- these genera and between species of As}>len- phologv, E. virginicum is very similar to E. ium, based upon sterile foliage alone. Clarifi- hjeUi Mantel. cation of resulting taxonomic problems awaits Equisetum marijlaiulicum was described in furthcT collection and study of existing speci- 1889 bv Fontaine. The aerial stem of this mens. Biological Skhies, \oi.. 14, \( Dakota Sandstone Floha 15

illustrating pirniuk- shapt- and disposition. "(X12:5). Fic. 7. 1. Glcicheniu dcliciilutu Ht-t-r. Portions of pinnae pinnule shape and disposition. BVU 1829. 2. GleinchenUi dclicatula Heer. Portions of pinnae illustrating rhizome illustrating shape and disposition of tub- (X.3). BYU 1829. .3. Equisftum lijclli Mantel. Portion of fragment illustrating characteristic branching. ers. (X.3). BYU 1809. 4. Gleicheniu dclicatula Heer. Pinna illustrating tubers with subtending bracts. (X3). BVU 18.30. .5. Equlsetum Itjclli Mantel. Portion of rhizome (X3). BYU 1810. 16 Bhk;ii\m ^()^ \(; Univkhsitv Sciknce Bui.lktin

Aspleiiitiiit (licksoiiianutu Ilet-r ing to give rise to two r;uhi; piniKie alternate, 35 or more long In 11 mm wide, lance Fi^s. 8-4; 11-3; 12-2; 15-4. mm olate; pinnules ovoid, up to 9 mm long by 3.5 1874 As])Ieniuin dicksoniamim Hcht— Flor. Foss. mm wide, alternate, attachment basal with Arct., V. 3, pt. 2, p. 31, PI. 1, Fig.s. 1 (exd. b, c), rounded sinuses, to nearly single point, apices laa, 2-4, 5 (excl. a, b). obtuse to rounded, margins entire; midvein 1899 As])leniuin (licksonininmi Ilccr, VVard- strong at point of origin, not extending to pin- dixiding; secondarx veins mosth' ob- U.S. Geol. Surv. 19tli .\iiii. Kept., pt. 2, p. 704, nule apex, to pinnule margin; fer- PI. 170, Fig. 1. scure, free, extending tile specimens imknown. 1950 As])lcniiiin .sp. brown— U.S. Cool. Surv. Prof. Paper 22 1 -D, p. 48, PI. 10, Fig. 4. Occurrence.—Known from the Dakota Sand- stone Formation near Westwater, Grand f'oun- Descrh'tion.— ICntiri' leaf unknown, at lea.st bi- t\', Utah. pinnate; ultimate pinnae alternate, up to 90 mm or more long bv 50 mm wide; pinnule.s lanceo- Repository.— Brigham Young University, Iiolo- late, entire to deeply cut, altcrnati', up to 40 t\pe; BYU 1806.' mm long bv 9 mm wide; attachment ba.sal to Dificvssio\. —Asjilcnium dakotensis is proposed .single point; midvein .strong at point of origin, rather delic;ite Asiilcniuni from the not extending to pinnule apex, dividing; .second- for a small, flora. It differs from A. dick.wnian- ary vein.s repeatedh' divided, free throughout; Westwater tini from the s;ime locality by being much fertile .specimen.s unknown. Heer smaller, both in pinnule and pinna size, and by OccuHRENCE.— Knowni from the Dakota Sand- pinnules being entire and rather strap-shaped, stone Formation near Hot Spring.s, South Da- rather than dentate or deeph cut and lance- kota, near Naturita, Montrose C'ountv, Colo- olate as are pinnules of A. dicksonwnum (Table rado, and near Westwater, Grand County, Utah. 1). A. dakotensis Rushforth is also much small- Repository.—Brigham Young University, BYU er than most other species of Asjilcniuni and 1811-1814. differs from m;uiv by Ii;iving pinnules with en-

tire margins. It is also rather unique in its bi- Discussion.—The author has elected to follow furcated petiole which gives rise to two rachi. Ward (1899) who collected an asplenioid fern This last characteristic coupled with ]iinnule from the Dakota Sandstone of the Black Hills shape and margination led the author to con- region and pliiced it in A. dicksonianuiu. How- sider this fern closely related to Knowltonella ever, Brown ( 1950), in describing ;i simihu' fern Berr\' (1911). However, examination of the type collected 1)V Stokes from the Dakota Sandstone specimens of KnowltoncUa indicates that the of Montrose (bounty, Colorado, noted that both branching habit of the two differs, as does the Colorado and the South Dakota specimens pinnule shape. Fertile specimens of neither appeared to be somewhat smaller and more A. dakotensis have been dis- delicate than typical specimens of A. dickson- Knowltonella nor covered, and for the present, the author prefers ianum. However, b;ised upon material in his assign this fern to the Aspleniaceae ;is As- possession. Brown (1950) was hesitant to name to j)Icniutn dakotensis, rather than to the Maton- a new species of this genus and so ascribed the iaceae imder KnoulfonclJa. material to AsjileiUuin sp. The observ;itions of Brown were accurate T;il)!e 1 for some specimens ol Asj)lciututi from Colo- rado. However, with the smaller forms men- Comparative M I leer am tioned bv Brown (1950) are inanv which are larger and resemble rather closely A. dirkson- kmuin. Westwater specimens of Asplenitini are also like A. dicksonianuin. although positive identity of this feni awaits collection o( specimens illustrating soral and spor;ingiaI characteristics.

Asplenitini dnkolcnsi.'i Hushfortli, sp. nov.

Kij^s. 11 -.5; 12-(i.

Descuii'tion. —Entire leaf unknown, bipinnate; petiole smooth, 1-2 mm in di;imeter, bifurcat- 17 ione Fi.oha Bioi.ocicAi. Sehiks. \()1.. 14, No. 3 Dakota Sands

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(M.-). BVU 18-7 Pinna fragment illustrating clmracteristic hranchuig - Kir. 8 1 C.U-ichcnia drlicatulu Hver. long, hnear pnmnk^. (XI 2 „,««>„,„•„„,./»», Rushforth. Pinna fragment illustratn,g Ltoni^un hrownii var n Pinna fragment illustrating pinna and p.nnule cbspos, k H Z- 3. Cleichcuia' dclicutula Heer. ' nltmiate pnmae drspos.t.on. X .2 \1 1828 4 Asplcninm ,licksouu.num Heer. Pinna illnstrating ) BVU chsposition. (X ChnlophkL pana Fontaine em, Berry. Pinna fragment illustrating pinnule BVU 81 1 5 eharaeter.sfc fragment . lustratmg 184V 6 comptonuwjoliu (Deb. and Ett.) Heer. Pinna BVU GlcichcnU, of dick.somanum Heer. Pinna fragment dlustratmg disposition l,ranel'ing:'(X1.2) BVU 1817. 7. Asplcmum ultimate pinnae. (X1.2). BVU 1812. 18 BiutaiAM VouNc Univehsity Science Bulletin

Family 1:)K:KS0NIACEAE C. bella Harris, which is similar to C. burejensis (Zalessky) Seward also is similar to C. Genus CONIOPTERIS Brongniart westwaterensis. C. bella is only separated from C. burejensis based on pinnule margination Coniopleris is probably the best known fos- and fine soral details, and C uestwaterensis sil genus of the fern family Dicksoniaceae. As this fern as from C. Imrejensis. C. listed bv Jongmans (1959) this genus contains differs from approximateh- 45 species and three varieties. westwaterensis also differs from both of these that it exhibits secondary pinnae which However, as discussed by Harris (1961) this ferns in are consistenth' larger, being up to 20 mm wide, genus is in need of revision, and fewer than 45 of C. burejensis and C. bella species are likely to be valid. whereas the pinnae are typically 10 mm wide.

Conioptcris uestwaterensis Rushforth, sp. nov. Family DIPTERIDACEAE Fig.s. 10-2; 11-4; 13-5. Dunker Description.—Entire leaf unknown, at least bi- Genus HAUSMANNIA divided into a sterile portion be- pinnate, some Dunker (1846) proposed this genus for ster- portion above; sterile pinnae low and fertile ile leaf fragments collected from the Wealden greater than lanceoiale, up to 20 mm wide by of Germany. These fragments were divided by alternate, 60 mm long; pinnules suboppositc to nearlv ecjual ditchotomies to form strap-shaped wide, lanceolate, up to 12 mm long b\' 3 mm leaf segments. Subsequent to this time, several strongly den- margins entire near pinna apex to other specimens of this genus have been col- generally tate further down the pinna, apices lected, and a rather adeejuate generic diagnosis attach- acute but range to somewhat obtuse, has evolved (Harris, 1961). ment basal to constricted basal, sinuses rounded (pinnules connected along rachis); midvein Ilatisuiannia rii^ida prominent; secondary veins mosth' obscure; fer- Newberry tile foliage consists of sterile pinnae below and ¥\gs. 9-2, 9-3; 13-4. fertile pinnae above; fertile portion of frond up 1895 llausmunnia rigida Newberrv— U.S. Geol. to 65 mm long or longer above highest sterile Surv. Mon. 27, p. .35, PI. 1, Figs. 2, .3', 5. pinna; fertile pinnules highK' reduced, alternate, connected along the rachis; venation ob- 1910 Newberri/ana rigida Berry—Jour. Geol., v. not observed. scure; sori, sporangia and spores 18, p. 254.

OccuHHENCE.— Known from the Dakota Sand- 1911 Neivbern/anu ri<^ida Hcri-y— Geol. Surv. stone Formation near Westwater, Grand Coun- New Jersey Bull. 3, p. 220. ty, Utah. Desciui'tion.— Entire leaf unknown, divided to Repository.—Brigham Young Universitv, holo- form tongue-shaped pinnules; pinnules typical- 180,3. type: BYU l\- 9 mm long b\- 3 mm \\k\v. apices rounded, margins entire, attachment basal with rounded Discussion.—This fern is known from several sinuses; midrib prominent, apparently not ex- specimens collected from the Westwater local- tending to pinnule apex; secondary veins ob- tv. A few of these are rather complete, illus- scure if present; fertile specimens unknown. trating fertile as well as sterile foliage. How- ever, even though fertile specimens have been OcciiRHENCE.— Known from near Westwater, collected none illustrate sori or sporangia. Ciriuul (bounty, Utah.

C. westwaterensis is similar in some respects Repository.- Brigham Young Universitv, BYU to several other species of Coniojiteris. It per- 1815. haps approaches most closely C. I>ttiejen.sis (Zalcssk\) Seward which is a common species Discussion.—The botanical affinity of llausnuin- from the Mid Jurassic of Asia and has also nid luis been in (juestion in the past. Some early

been collected from Alaska and Europe. C. workers considered this plant to be ;i hepatic. westwaterensis differs from C. burejcnsis (Zales- Newberry (1885) said in this connection: skv) Scuard in having less crowded pinnae and

1 It'll think it possible it was a bv having the fertile portion of the foliage li;i\c bffii to liiUhtT kiiul of hepiitic-, a Marchanliu. lor cx- separated from the sterile portion, often oc- amplf, lifted from its cr<-i-piiig (.oiulitioii into curring as much reduced pinnae apicalK' to an iiulfpt-nilent and erect phint, trained and sterile pinnae. disciplined into syininetr\ by the occult in- Biol

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Ay;y -^^^^ i' ^

Fic. 9. 1. Astradopteris colurudicu (Brown) Reveal, Tidwell, and Ruslifortli. Near apical pinna fragment. (X1.2). BYU 1839. 2. Ilansmannia rigidci Newbern,'. Near apical pinna fragment illustrating tongue-sliaped pin-

nules. (X1.2). BYU 1815. .3. llausmunniu rigidu Newberry. Enlargement of Fig. 2. (X3). BYU 181.5. 4. Gleichcnia delictitula Ilccr. Piima fragment illustrating disposition of ultimate pinnae. (X1.2). BYU 1826. 5. Cladopldchis crenuta Fontaine. Pinnule fragment illustrating venation. (X3). BYU 1841. 6. Gleichcnia

comptoniucfolia (Deb. and Ett. ) Heer. Pinna fragment illustrating disposition of ultimate pinnae. (XI. 2). BYU 1816. 7. Astmlnptcris roloriidica (Brown) Rexcal, I'iducll, and Rushforth. Pinnae fragment. (X1.2). BYU 1840. 20 BmcuAM YoiNc; University Science Bulletin

fluencc uhicli lias given siuli grace and ex- tiiat tiie divisions of the leaf are not dichoto- actness to the foliage of fenis, lycopods, and mous, and are more regular in the former. some conifers.

Ihtusiiumnia Later studies have shown tluit Family GLEICHENIACEAE is ;i fern of the family Dipteridaceae ( Richter, 1906), although fertile speeiiuens demonstrat- Ceims GLEICHENIA Smith soral characteristics are still unknown. The in<4 Berry (1924) discussed the usage of Glei- Westwater llausmannki doc>s not shed any light cJienites and Glciclicnia. In this paper, Berry problem since fertile specimens are un- on this rather stronglv supported the use of Glcichcnia known. rather than CAcichenitcs and stated in this re- first treated in America by llmismannui was gard, "It is surelv nothing but a mental illusion Newberry (1S85) who described //. rigida. This to imagine that the iLse of an objectionable species was proposed for several plant fragments tenn like Clciclicnitcs indicates a conservation the Clays of New Jersey. collected from Amboy of judgment." However, Seward ( 1927 ) dis-

Berry ( I9I0) later transferred llausmannia agreed with Berry and used the name Gleichen- rigida to the new genus Newbern/ana without itcs for placement of several Greenland species. description, discussion, or illustration, and Seward stated that in his opinion it is a sound hence, his reasons for this move were unknown. practice to use an ite.s ending for a generic However, Berry (1911, p. 220-221) further name when "either our ignorance or the occur- discussed Haiisni'innia rigida and stated that rence of some character in which a fossil species Newberry had referred this plant to the Hep- differs from anv existing type" may apply to an aticae, and due to this: extinct taxon. Harris (1961) pointed out that the practice It is, obviously, not related to the genus llaiis- of using genera with an ites ending is being mitnnia, Dunker, which has been definitelv dropped bv most paleobotanists in recent times. proven to be a fern genus of the familv Dip- Harris, however, rightfully suggested that when teridaceae, so that I have selected to propose a exists as to whether a fossil taxon may . question a new generic name. . . with ecjual validity be placed with more than However, Berry gave no reasons why this plant one living taxa, a different generic name for should not be considered to be related to Haus- the fossil must be used. nuinnia, and further went on to say that indeed It is my opinion that it is sound to use the it is probably a fern. It is the opinion of the ites ending under certain circumstances when present author that Newberry was justified in evidence of the sameness of a fossil and living placing his material in llausmannia as a new genus is suggested but not definite. However, species since this material agrees rather well the use of this ending merely because of the with the generic diagnosis of JIausiminniu. Fur- older age of a fossil is not valid. When there ther collections and reevaluation of Newberry's is little or no (juestion that a fossill may be material should, however, be undertaken. placed in an extant genus, it is both acceptable and desirable to use the modem generic name //. rigida Newberry ( 1895, p. 35, PI. 1, Figs. 2, 3, 5.) is probably most closely related to the for its placement. Westwater llausmannia. This species lacks di- The fossil section of the genus Glcichcnia cholomous leal ilivisions, and more regular divi- is in urgent need of revision and reevaluation. sions of the frond-forming tongue-shaped seg- Many species are undoubtedlv svnonyms and ments is also found in the Westwater species. could be more correctly placed. In addition, As discussed by Nevvberr\- (1895), the midrib this genus afford.s an excellent opportunitv to in //. rigida gives rise to numerous fine secon- follow the cNolution and migration of a tern dary veins, although his illustrations do not dem- genus from Paleozoic to recent times, and its onstrate this feature. These secondarv veins study could contribute greatly to botanical cannot be observed in the s[)ecimen from West- knowledge. water.

//. dicliotonui Dunker is a common species Glcichcnia coniptoniaefolia ( Deb. and of this genus from the Jurassic and Lower Cre- Ett.) Heer taceous of Europe. This species usually has Fig.s. 8-6; lO-I, 10-6; I2-I, 12-3; 13-1. been well described and, as mentioned by Neic- 13-3; 14-3; 20;i. berry (1895), is closely related to llausmannia rigida. II. rigida differs from //. dichotoma in 1859 Didt/tnosarus coniptoniaefolia Deb. and Bi()L()(:ic:al Skuies, Vol. 14, No. 3 Oakoia Sandsionk I'i.oha 21

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Fig. 10. 1. Gleichenia comptoniuefoliu (Del), and Ett. ) Heer. Pinna fragment illustratiiif; disposition of ultimate pinnae. (XI. 2). BYU 1821. 2. Conioptehs wcstwatercnsis Rushfortli, sp. nov. Pinna fragment illustrating dis-

|X>siti()n of fertile pinnae. (X1.2). Paratope BYU 1804. .3. Matonklium hrownii Hirsliforth. Pinna fragment illustrating pinnule disposition. Tiiis fragment is .somewhat atypieal and resembles some speeies of Sclenocar-

/)i/.s- Selienk. (XI. 2). BYU 1833. 4. Gleiclwnki dcUcntula Ileer. Pinna fragment illustrating ultimate pinnae ilis|«>sition and pinnule shape. (X1.6). BYU 1829. .5. Miitonklium hrownii Hushforth. Pinna fragments and fiddle head. (X1.2). BYU 1834. 6. Glciclwnia coviptomacjolm (Deb. and Ett.) Heer. Pinna fragment illustrating ultimate pinnae disposition and pinnule shape. (XI. 2). BYU 1822. )

Bhicham Vounc Univehsity Science Bulletin

Ett.-Denksch. K. Akad. Wiss. Wicn., 17, PI. 1, -U.S. Geol. Surv. Mon. 15, p. 119, PI. 21, Fig. Figs. 1-5. 11.

1868 Gleichenia gieseckiarui Heer, pro partt-— 1889 Gleichenia zippei ( Corda ) Heer, Fontaine

Flor. Foss. Arct., v. 1, p. 78, Pis. 43, Figs, la, -in Ward, U.S. Geol. Surv. 19th Ann. Kept., p. 2a, 3a; 44, Figs. 2-3. 664, PI. 162, Fig. 9. 1868 Pecopteris borealis Brong., Heer, pro parte 1927 Gleichenites gieseckiana (Heer) Seward—

-Flor. Foss. Arct., v. 1, p. 81, PI. 44, Fig. 5. Phil. Trans. Rov. Soc. London B, v. 215, p. 69, PI. 5, Figs. 1-4,' 6-14, 16, 17; PI. 10, Fig. 96; PI. 1868 Gleichenia zippei (Corda) Heer-Flor. 12, Fig. 118; text-Fig. 2. Foss. Arct., V. 1, p. 79, PI. 43, Fig. 4. 1950 Gleichenia kurriana Heer, Brown-U.S. 1868 Pecopteris arctica Heer, pro parte— Flor. Geol. Surv. Prot. Paper 221-D, p. 48, PI. 10, Foss. Arct., V. 1, p. 80, PI. 43, Fig. 5. Figs. 5-6. 1868 Pecopteris rinkiiina Heer— Flor. Foss. 1956 Gleichenia gieseckiaiui (Heer) Seward, Arct., V. 1, p. 81, PI. 44, Fig. 4. Bell-Geol. Surv. Can. Mem. 28>5, p. 61, PI. 17,

1874 Gleichenia longipennis Heer, pro parte— Figs. 1, 2; PI. 18, Fig. 5. Flor. Foss. Arct., v. 3, pt. 2, p. 46, PI. 6, Figs. leal at least bi- 4-6. Deschiption. —Entire unknown,

pinnate; rachis stout, slightly greater than 1 nun 1874 Gleichenia tliulensls Heer— Flor. Foss. in diameter, smooth, appearing dichotomously V. Pis. Fig. 9b; Arct., 3. pt. 2, p. 4S, 5, 10, branched (although not truly dichotomizing Fig. 18. [Bower, 1926] ), with a bud in the axil of dichot-

1874 Gleichenia comptoniaejolia ( Deb. and Ett. omy; ultimate pinnae alternate, occasionally sub- Heer, pro parte— Flor. Foss. Arct., v. 3, pt. 2, opposite, nearly linear, approximately 80 mm p. 49, PI. 11, Figs. 1-2. long by 10 mm wide; pinnules arising obtusely from rachis, narrow eliptic to deltoid near pinna 1874 Gleichenia gracilis, Heer, pro parte— Flor. apex, oriented slightly toward pinna apex, rang- Foss. Arct., V. 3, pt. 2, p. 52, PI. 10, Fig. 5. ing from 1 mm wide by 2 mm long at apex of 1874 Gleichenia gieseckiana Heer— Flor. Foss. pinna to 3 mm wide by 6 mm long near pinna

Arct., V. 3, pt. 2, p. 43, Pis. .3, Id, 8; 7, Fig. 1. base, margins entire, apices obtuse; major vein strong at point of origin, not extending to pin- 1874 Gleichenia nauckhoffi Heer— Flor. Foss. nule apex; secondary veins numerous, dividing, Arct., V. 3, pt. 2, PI. 25. Fig. 4. extending to pinnule margins; sori commonly 1874 Gleichenia zippei (Corda) Heer, pro three per pinnule, round; sporangia and spores parte-Flor. Foss. Arct., v. 3, pt. 2, p. 44, 90, 97, unknown. Pis. 4-7; 25, Figs. 1-3. OccuRKENCE.— Near Westwater, Grand County, 1874 Pecopteris hijperborea Foss. Heer— Flor. Utah, and Rabbit Valley, Grand County, Utah. Arct., V. 3., pt. 2, p. 41. Rei'ositoky.— Brigham Young University, BYU 1874 Gleichenia kurriana Heer, Lescjuereux- 1816-1825, 1853a^

U.S. Geol. Surv. Terr. Kept., v. 6, p. 47, PI. 1, Discussion.—As may be deduced from the syn- Figs. 5, 5b, .5c. onomv for G. comptoniacfolia, Heer proposed 1882 Gleichenia geiseckiana Heer— Flor. Foss. many species of Gleichenia in the late ISOOs Arct., V. 6, pt. 2, p. 6. PI. 2, Fig. 9. based upon material collected from several Greenland localities. Heer treated any fonn that 1882 Gleichenia zijipei ( Corda ) Heer- Flor. illustrated any variation whatsoever as a new Foss. Arct., V. 6. pt. 2, p. 7, PI. 3, Fig. 2. species. Seward (1927) reviewed the work of 1882 Gleichenia longipennis Heer— Flor. Foss. Heer and studied many of his original collec- Arct., V. 6, pt. 2, p. 7, PI. 2, Fig. 5. tions. Based upon his studies, Seward concluded 1882 Gleichenia comjHoniaefolia (Deb. and that many of the species proposed by Heer merely represent variations of a few species. Ett.) Heer-Flor. Foss. Arct., v. 6, pt. 2, p. 8, 36, PI. 46, Fig. 25. In discussing the synonomy tor C;;. giesecki-

ana, into which Seward ( 1927 ) lumped many of 1883 Gleichenia norclenskioldi Heer, Lesquereux Heer's ( I86S, 1S74, 1872) species, Seward men- -U.S. Geol. Surv. of Terr. Repts., v. 26, 8, p. tions two names which predate G. gieseckiana. PI. 1, Figs. 1, la. The first of these Pecopteris borealis Brongniart, 1889 Gleichenia nordenskioldi Heer, Fontaine was applied by Brongniart (1828). Seward ex- Biological Series, Vol. 14, No. 3 Dakot.\ Samwionk Floha 23

Fig. U. 1. Cladophlebis constricta Fontaine em. Bern'. Pinnules illustrating deeply lobed margins. (X3). BYU 1941. 2. Clmlophlehis puna Fontaine em. Berr\'. Pinna fragment illustrating pinnule shape and venation. (X3). BYU 1842. 3. Asplenium dichsnnkmum Heer. Pinna fragment illustrating shape and venation of pinnules. (X3). BYU 1814. 4. Coniopteris iLcstwtiteremis Rushforth, sp. nov. Portion of pinna illustrating pinnule shape and margination. (X3). Paratype; BYU 1805. 5. Aiplcniurn Jakotcnsis Rushforth, sp. nov. Portion of frond illustration pinnae and pinnule disposition. (X3). Holotype: BYU 1806. .

Bni(;nAM ^ OuNo Univehsitv Science Bulletin aniined the type specimen of this fern and con- discussed gieicheniaceous ferns. He placed a cluded that it was not possible to detennine if it specimen of Glek'henia in G. zippei (Corda) is synononioiis with G.

( 1859) is also an older name which has been ap- ervosa Velen. plied to C. gk'seckiuno Heer. Heer recombined Brown ( 1950) reported on gleicheniou.s fos- D. comptonkiefoliiis Deb. and Ett. as GlwcJienia sils from the Dakota Sandstone of southwestern gieseckkimi since the fonner is undoubtedly Colorado. He placed these ferns in Gleichenia gieicheniaceous. Seward (1927) placed C comp- kurriana following Lescjuereux (1874). Recent toniaefoUa ( Deb. and Ett. ) Heer in synonomy examination of these ferns by the present author with G. gieseckkimi Heer. Seward did not use the indicates that they are indeed very similar to oldest specific epithat l>ecause "evidence of iden- those figured by Les(|uereux (1874) as G. kur- tity is not convincing." However, Seward m the rianu which are correctly placed with G. comp- same pajx^r noted that some of the feni.s Heer toiiidcfolia. had placed in G. coniptonwefoUu were.without Bell (1956) described Gleichenites gieseck- question, referable to G. gwseckkma. Further- ianus Heer em. Seward from several localities in more, the illustrations of Debey and Ettinghau- western Canada. These ferns agree with the sen ( 18^59) indicate that it is very likely that the Westwatcr Gleichenia in all respects except that ferns referred by them to Didtjmosarus comp- the pinnules on the Canadian specimens are often toniaejolius are identical to G. gieseckkina. In somewhat reflexed. Again, these ferns are correct- view of this, it is preferable to use the oldest Iv classified as Gleichenia cornptoniaefolia ( Deb. specific epithat, and therefore the most accept- and Ett.) Heer. able binomial applied to G. gieseckiana of Heer, Seward ( 1927 ) discussed Gleichenites nor- is G. comptoniaefolki ( Deb. and Ett. ) Heer. detiskioldi Heer, and retained it as a valid species Lesquereux (1874) figured a gleichenious wliich differs from G. comptoniacjolia by having fern from the Dakota Group of Kansas. These long pinna of uniform breadth and pinnules with fenis agree in ever\' detail with Glek'henia somewhat more frecjuent veins. Soriation in the comptoniaefolki ( Deb. and Ett. ) Heer, but were two species is similar, but Seward noted that G. placed in Glewhenki kurrkiiui Heer by Les(|uer- nordenskioldi has more sori per pinnule. All of eux. However, it is important to note that Les- these features are extremely variable, and speci- (juereux considered the Dakota ferns closely al- mens from Westwater may be found intermediate lied to Duit/mososorus comptoniifoJius Deb. and to all of these species-differentiating character- Ett., and in fact noted that in pinnule shape, istics. However, the author has elected not to disposition, and venation, "our American species make any changes in classical usage of G. nor- or variety would be more closely related to denskioldi and G. comptoniacjolia until further ." Dklymososonis comptoniifoUus Deb. and Ett. . review.

Les(|uereux ( 188.3) again treated gliechenious Gleicheneous fossils from the Westvvater lo-

ferns in his work on the Cretaceous and Tertiary cality are similar to those figured by Heer ( 1868, of the Western Territories. Pinna fragments of 1874, 1882), Seward (1927)"; and others. Similar Gleichenia from the Dakota Croup were placed problems of classification have been encoimtered by Les(|uereux in G. nordcn.skwldi. However, as with these ferns in that they exhibit rather wide

noted bv Seward ( 1927 ) the feni examined by morphological variation. However, the present Lesquereux agrees in all respects with G. giescck- author has elected to be more conservative than iana (G. comptoniacjolia) Heer and Seward, and many fossils which exhibit Fontaine (1889) discussed Glewhemu nord- some degree of variation are nonetheless placed enskioJdl Heer as known from the Fotamac in G. comptoniacjolia (Deb. and Ett.) Heer.

Croup. However, this fern agrees with G. comp- This species of fem is the most common ele-

tonkicfolki closely, and should be placed in this ment from the Westwater area and it is present species. in such profusion that at certain outcroppings In discussing Lower Cretaceous plants from one can scarceh' break open a rock without ex-

the Black Hills region, Fontaine (1899) again posing at least tme beautiful specimen. It is in- Biological Series, Vol. 14, No. 3 Dakoia Sandstone Flora 25

• f-:-:-,

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ultimate pinnae and pinnules. Fig. 12. 1. Gleicheniu compUmuwfnlia (Deb. and Ett.) Heer. Pinna illustrating disposition of ultimate pin- (X1.2). BVU 1818. 2. A.splenium dicksimianum Heer. Pinna fragment illustrating Heer. Pinna illustrating ultimate pinnae. (X1.2). BYU 1812. .3. Gleicheniu complnniaefolia (Deb. and Ett.) characteristic shape nae and pinnules. (X1.2). BYU 1819. 4. Sniix newhern/ana Hollick. Leaf demonstrating shape and margin- and margination. (X1.2). BYU 18.51. 5. Ilex .serrtilus Rushforth, sp. nov. Leaf illustrating nov. Pinna illustrating shape and ation. (X1.2). Paratype; BYU 1802. 6. Aspletiium dakotensis Rushforth, sp. disposition of ultimate pinnae and pinnules. (X1.2). Holotype: BYU 1806. 26 Bhic.ham Vou.nc Univeksitv Science Bulletin

Fic. 13. 1. Cleichcnm comptoniacfolut (Deb. and Ett. ) Heer. Pinna frajrment iUustratint; soral disposition. (.X.3).

BYU 1823. 2. Clciclicniii comptonuwfolia { Del), and Ett. ) Hccr. I'innae frapnents u ith pinnnlcs exhibiting illnstrating venation. (X3). BVU 1824. 3. Cleichcnm compltonUicfoliu ( Deb. and Ett.) Heer. Pinna fragment characteristic branching. (X3). BVU 182.5. 4. Hausrrmnnia rigida Newberry. Pinnae fragment demonstrating tongue-shaped pinnules. (X3). BYU 181.5. Biological Series, Vol. 14, No. .3 D.\kot.\ Sandstone Flora 27 teresting to note that gleicheniaceous ferns are the pinnule shape, which as described by Heer also most common from the Greenland Cre- is round in the Greenland specimens, but tends taceous flora. Indeed, as mentioned by Seward, toward ovate with an angular apex in the West- ". . . part of the country was occupied b\' a water material. In the oblong pinnule shape, vertiable Gleichenietum." the Utah material resembles Gleichenia micromera Heer, a contemporaneous species with G. delicatula. However, the westwater material is Gleichenia delicatula dissimilar to G. micromera in that the pinnules Figs. 7-1, 7-2, 7-4; 8-1, 8-3; 9-4; 10-4. in the latter are not attached to the rachis with their entire base, whereas, they are broadly at- 1874 Gleichenia delicatula Heer— Flor. Foss. tached and with rounded sinuses in the Utah Arct., V. 3, pt. 2, p. .54, PI, 9, Figs, lie, llf; PI. Gleichenia. 10, Figs, 16, 17, Velenocsky ( 1888 ) discussed and figured 1882 Gleichenia delicatula Heer— Flor. Foss. two specimens of Gleichenia delicatula. These Arct., V, 6, pt, 2, p. 9. specimens agree in all details with Heer's (1874) specimens, and differ from the West- 1888 Gleichenia delicatula Heer, Velenovsky— water specimens only in pinnule shape. Pinna Abh. K. Bohm, Ges. der Wiss., Math.-Natw., morphology and size of Velenovsky's specimens ser. 7, V. 1, p. 7, PI. 3, Figs. 12-14. is identical to that of the Utah Gleichenia. 1910 Gleichenites delicatula (Heer) Seward— Seward (1910) used the generic name Glei- Fossil Plants: v. 2, p. 3.54, Fig. 262b. chenites for the placement of Gleichenia delicatula. He made no changes in description, and 1919 Gleichenia delicatula Heer, Berry— U.S. figured a specimen of Gleichenia delicatula Geol. Sury. Prof, Pap. 112, p, 54. which appears very close to those of Heer Description.—Entire leaf unknown, at least bi- (1874), Velenovsky (1883), and the material pinnate; petiole less than 1 mm in diameter, from Utah. Again, the pinnule shape varies be- smooth, appearing dichotomously branched with tween Seward's specimens and those from Utah, pinnae a bud in the axis of dichotomy; ultimate Berry (1919) reported on Gleichenia deli- alternate, linear, up to 15 mm long by 2 mm catula collected from the Tuscaloosa Formation wide; pinnules tending toward oyate but with of Fayette County, Alabama, This fern was truncated edge toward pinna apex, oriented placed with G. delicatula based upon the pin- slighth' tovyard pinna apex, approximately 1 mm nules being united to the rachis along their en- 1.2 wide, attachment basal with long by mm tire base. Berry further mentioned that Fon- rounded sinuses, margins entire, apices acute to taine (1895) had wrongly placed pinna frag- obtuse, directed apically; primary yein diyiding ments collected from the Raritan Fomiation in immediately upon arising from rachis; secondary G. niicroniera. Berry placed these in G. deli- five ulti- yeins dividing, giving rise to three to catula based upon their completely basal at- mate veinlets which extend to pinnule margin; tachment. Again, this fern is similar in size to fertile specimens unknown. the Westwater fern, but pimiule shape differs Occurrence,—Near Westwater, Grand County, somewhat. in pinnule the West- Utah, and Rabbit Valley, Grand County, Utah Even though shape water Gleichenia differs somewhat from other Repository.— Brigham Young University, BYU small pinnuled species of this genus, for the

1826-17.30. present time the author has elected to place it

with G.delicatula Heer, This placement is based Discussion.-G/eic/ierii'a delicatula was proposed upon similar pinna morphology and pinnule size by Heer in 1874 for placement of several pinna and disposition. fragments collected from the Greenland Cre- taceous. This fern was originally described by Heer (1874) as follows: Family MATONIACEAE Gl. fronde gracillima, (lichotoma, bipinnata, Genus MATOIIDIUM Schenk racchi tennuissima, pinnis approximatis, pat- entibus. linearibiis, pinnvilis miiiiitissinms, ro- Schenk (1871) was the first to use the gen- tundatus. eric name Matonidium for placement of fossil This description, and the illustrations of ferns exhibiting similar characteristics as the ex- Heer agree rather well with the Westwater tant Matonia pectinata \\. Br. Fossil ferns placed specimens. One readily apparent difference is in this genus are morphologically very similar 28 l$Hi(;nAM ^ouNG Univebsitv Science Bulletin

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Fk;. 11. 1. Mdloiiidiiiiii hrouiiii Hiishtditli. I'iMiia iraKmi'iil illu>tiating \t'ii.iti(>ii .iiid soriatioii <>l piniuik'S. (.\1.2).

BVl' I5fi3. 2. Mdtdiiidium hrownii Busliiortli. Kiihirncineiit of Fijj. 1. (.\2.7). ti\'V 156;5. 3. Clcichcnia comptunmefolia (Del), and Ktt.) Heer. I'iniia fraf^moiil illiistiatini^ ultimati- pinnae disijosition. (XI. 2). BYU 1820. 4. Astrdloptcris coloraclUu Heveal, Tidwfll, and Huslifortli. Pinna fragment illustrating pinnule attaeh- ment and shape. (\1.2). BVU 1838. 5. Coiiiopteris urstwiitcrcn.'iK Bushforth, sp. nov. .Sterile and fertile pinnae. (.\1.2). Holotvpe: BWJ 1803. (i. .\ngiosperni leaf base possihK...helduging to h'icus sp. (X1.2). BYU 1847. Biological Series, Vol. 14, Ni). 3 Dakota Sandstone Flora 29

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Fic. 15. 1. Miitanidiuni brouuii Rusliforth. Pinna apt'x dfmunstratini^ disposition of pinnules. ( -XI. 2). BYU 1832 2. Matonidiuni hroiinii Hiishforth. Pinna fragment demonstrating venation and soriation. (X2.8). Para- type: BYU 15.59. 3. Malonidium (?) lanicipinmiliim linshforth. Pinna transfer illustrating pinnule shape and disposition. (X1..5). Holotype: BVl' 1569. 4. Asiileiiiunt dicksonianunt Heer. Pinna transfer illustrating pinnule disposition and venation. (X2.4). BYU 1813. 5. Eqiii.\ctitin li/clli Mantel. Portion of rhizome with two tuhers attaehed. (X1.2).B^'U 1807. 6. Mutimidium hrouitii var. magniiniiiiuluni Rushforth. Pinna fragment showing long, linear pinnules. (X1.2). BYU 18.36. 7. Satix netcherr\janu Hollick. Leaf demonstrating characteristic shape and size. (XI. 2). BYU 18.50. 8. Eucahjptus dakotenxis Lesquereu.x. Leaf of characteristic size and shape. (X1.2). BYU 1848. 9. Equisctum Itjclli Mantel. Rhizome transfer with attached tubers. (X1.2). BYU 1808. 30 HniciiAM VouNc. University Science Bulletin and often essentiallv identical to modem maton- feature is easily discerned in the specimens ex- iaceous ferns. amined by the present author and may be the

Matonidium is often difficult to differentiate reason that venation in this species is poorly pre- from other members of the Matoniaceae, par- served. Berry also described the pinnules of this ticularly PhU'l)Oj)teris. However, Matonidium is species as having revolute margins. While this separated from PliIcl)Oj)trri.s liased upon the pre- is possible, it is likely that this appearance is sence of an indusium in the loniier. This differ- due to the coriaceous nature of the pinnules. ence is often very difficult to determine even in This feature is (juite unicjue within the Maton- fertile specimens, and is impossible to ascertain iaceae. in sterile material. Fiirtliermore. to compound This species also differs from other species this problem, Matonidium and Phlebojiteris are of Matonidium in the large number of pinnae often very similar in venation. Further work bome on a single frond. In addition, the open dealing with this family will likely show that the venation pattern is somewhat uncommon al- distinction I)etween these two genera is unnat- though not unprecedented within the Maton- ural and that ferns variouslv placed in the two should be classified under a single genus. Matonidium brownii Rushforth

Matonidium americanum Berry em. Rushforth FiK.s. 10-3, 10-5; 14-1, 14-2; 1.5-1; 17; 18-2; 18-1, 18-3. 18-6. Kij;s. 18-:5. 18-4; 19-.5. 1950 Matonidium americanum Berry, Brown— 1916 Matonidium althausii (Dunker) Ward, U.S. Geol. Surv. Prof. Paper 221-D. p. 48, PI. 10. Cockerell-J. Wash. Acad. Sci., v. 6, p. Ill, Fig. Figs. 7-8. 2. 1970 Matonidium hrounii Rishforth-BYU Geol. 1916 Cijcadospadix sp. Cockerell— J. Wash. Acad. Stud. 16(3), p. 9, PI. 1, Fig. 2; Pis. 2-5; PI. 6, Sci., V. '6, p. 111. Fig. 1. Fig. 2; PI. 8. Fig. 2; PI. 11, Fig. 1. 1918 Matonidium americanum Berrv— Bull. Tor- Description.- Petiole stout, between 5 mm ;ind rey Bot. Club, v. 46, p. 287, Fig. 2. JPls. 12, Figs. 15 mm in diameter, divided apicallv into 1-12; 13, .3-6. two short arms; pinnae four to ten or more 1970 Matonidium americanum Berrv, Rushforth per frond, lanceolate, up to 14 cm in length,

Geol. Stud., v. pt. .3, 25, PI. 8, Fig. -BYU 16, p. piniKi rachis between 0.5 mm and 2..5 mm in I; PI. 13, Fig. 2. di;uneter; pinnules opposite to alternate, basal in attachment, decurrent but free from each other, Description.—Fronds pedate, petiole stout, to up to 30 mm in length by 9 mm in width, arise approximately 1 cm in diameter, furrowed (?), nearly to the rachis dividing to form a collar from which pinnae are perpendicular but become oriented toward pinna apex; margins entire; bom; pinnae lanceolate to linear, 20 cm to 25 ;ipices rounded; midrib prominent to pinnule cm long, up to 39 per frond; pinnules inserted apex; secondary veins arise from midrib, divide near the upper margin of the rachis, up to 9 mm to form costal ;ieroles; tertiary veins arise from long by 3 mm wide, coriaceous, more or less falcate; apices rounded; margins entire or revo- costal areole and other secondary veins and anastomose in tlic pinnule lamiuii; costal areoles lute; midvein prominent to near the pinnule absent in fertile specimens; secondarv veins apex; secondary veins prominent, bifurcate ap- fomi an arch above placenta; tertiarv veins arise proximately one-half to two-tliirds of the way to from this arch extend to the pLiccnta; upper margin, remain tree; sori biseriate, round (ex- ;md epi(lcrm;il cells more or less isodi;mietric, aver- cept where crowded), decrease in size toward age 45u in diameter; lower epidemial cells un- pinnule apex, indusiate; indusium peltate; sporangia and spores unknown. known; sori round, biseriate 1.7 mm to 2.9 mm in diameter, with a single ring of from ten to OccuRRENCK.—Near Cutthroat Clulch and Hoven- fourteen sponuigia; indusi;i present, peltate, per- weep Canyon west of Dolores, Montrose Coun- sistent; sp()r;ingi;i cuneatt- sessile or nearly so; ty, Colorado, and Rabbit Vallev, Grand County, iinnuli obli(|ue; spores trilete, laesurae three Utah. (jUiirters of the distance to the efjuator, rounded- Repository.—Brigham Young University, BYU tri;ingular, 57u in diameter, with a weak margo. 1572, 1831. Occurrence.— Known from ne;ir N;iturita, Mon- Discu.ssioM.-As mentioned by Berry (1919) the trose ('ounty, Colorado, and nc;ir W'estwatcr, pinnules of M. anwricanum are coriaceous. This CJnmd (>ountv, Utah. BioLocK.AL Skries, \()l. 14. No, 3 Dakota Sandstone Flora 31

'"^ (M-2) BVU 1843 ^ Ilex sILJh T'^1 ""^'- ^"p.^"'*^-^'', AnRuIar-ovate leaf with typical serrate to Sm spinose Lr^in. (XI 2) Holot pe"' BYU 1801. 3 Ficus daphno<,cno,dc., (Heer) Berry. Cuneate leaf hase exhibitini- cuneate base XI 1845 "'''"-g-."'"'" Berry. f^^^' («-), Leaf e.xhibiting drip point, sbape and mar.ination. 5 Ji'X X ) BYU IH^ neer. rortion ot leal exhibiting characteristic- venation. (.\]..5). BVU 1849. Science Bulletin 32 Bhicham Young University

Hki'ositohy.— Brigham Young University, holo- Dlscussion.—M. hrounii var. m

placed this feni in ^^ . amcri- However, Brown rower than those of var. hrownii. In addition, Berr\'. The present author examined the canum the disposition of the pinnules on the rachis Ls collected from this locality and con- specimens farther ;ipart in var. nuiiinijiinntilum. cluded that these specimens should not be placed As mentioned by RusMorth ( 1970 ) , this fern with M. (imericamtm. Due to this. M. hrounii inav be distinguished from other matoniaceous Hushforth ( 1970) was proposed. ferns due to its large pinna and pinnule size. differs from M. americamtm by M. I)ruunii However, long, linear pinnules are not unknown larger; by having fewer piniuie per being much within the Nlatoniaceae. Bell (1956) reported frond; bv having anastomosing rather than open on Phk'hopteris (?) elongata for sterile pinna (Hushforth, 1970). venation; and in soriation fragments from the Cretaceous of Canada, and M. hroumi differs from a related species, M. P. imiensteri also exhibits long linear pinnules. uicstwri Krasser, by having anastomosing rather than open venation; by being larger in overall Miitonichuin lanci])innuhini Rushforth size; and bv having more numerous sori. M. (?) differs from M. (iltlmusii (bunker) Ward Figs. 1.5-:3; 18-1; 19-2. hrotinii in piniuile shape and habit, ;uid in ven- 1970 Matonidiurn (?) Iancii>innuhini Rushforth ation. Soriation of M. althausii also differs from Studies, v. 16 pt. 24, Pi. 7, Fig. M. hrotinii in that the fomier exhibits sori bom BYU C;col. 3, p. 1; PI. 9; PI. 10; PI. 13, Fig. I. to the extreme pinnule apex decreasing in size as the apex is approached. Description.—Entire leaf unknown; pinnules In venation M. hrounii is similar to Phleh- sub-opposite, linear-lanceolate, 60 mm long by

0})teris ( Schenk ) Schenk. 12 mm wide, essentiallv perpendicular to rachis except at extreme pinnule apex, attachment basal with rounded sinuses; midvein prominent to Matonidiurn hroicnii var. nuignipinnulum pinnule apex; secondary veins prominent, form- Rushforth ing costal areoles, anastomosing above c-ostal of Kigs. 1.5-6; 18-2; 2Ib. areoles except near laminar edge, composed tracheids with annular, helical and reticulate 1970 Matonidiurn hroicnii var. magnipinmdum wall thickenings; guard cells 4(hi long by 15u Geol. Stud., v. pt. 3, 11, Uushforth-BYU 16, p. wide, leaving stomata lOu wide; fertile speci- Pi. 6, Fig. 1; PI. 7, Fig. 2; PI. 14, Fig. 2. mens unknown. Dk.scriptiox.— Entire leaf unknown, pedate; pin- Occurrence.— Dakota Sandstone Fonnation near nae observed up to 30 cm long, longer in Westwater, Grand ("ounty, Ut;di. growth, lanceolate; rachis up to 1.5 mm in di- Repository.— Brigham Young University, holo- ameter; pinnules opposite to alternate, basal in " paratype: BYU 1579. attachment, often decurrent, free, up to 60 mm type: BYU 1569;

in length bv 6 mm in width, arise at ^X) but be- this fern DiscussioN.-Rushforth ( 1070 ) phiced oriented toward pinna apex; margins en- come in the Nlatoniaceae based upon pinnule venation prominent to pin- tire; apices rounded; midvein and shape. However, the assignment of this fern veins prominent, forming nule apex; secondary to this familv must remain provisional until fer- costal areoles in sterile pinnules, progressing di- tile specimens are collected. rectly to phicenta in fertile tonus; tertiary veins One interesting fact concerning this feni is often anastomose except in pinnule tips; lower that it seems to be somewluit intermediate in epidermal cells large (up to 95u by 45u), lobed; venation pattern betwi'cn wiil-di'tiiu-d members fertile specimens not observed. of the Matoniaceae and another WesU\'ater feni,

Occurrence.— Known from the Dakota Sandstone .\.stralo))t('ri.s ( Polypodiaceae ) . This latter genus Fomiation ne;ir Westwater, (irand C-ounty, Utah. illustrates \cnation which would tend to align

Reposiiohy.— Ikigham Young Universitv, holo- it with the \hitoniacc;ie, but sond characteristics ' type; BYU 1566; BYU 18.36. align it witli the Polypodiaceae. Further at- BioLocicAL Sehiks, \'()i.. 14, No. 3 Dakoia Sandstone Floha 33

Fig. 17. Mahmidium hroicnii Rushforth. Portion of a frond illustratinsr pinnae and pinnule di.sposition (.\12) f i Holotype: BYU 1557. Science Bulletin 34 Bhigham Young Univehsitv

Smithsonian tempts to locate more specimens of M. l(mci])in- with extant feni workers from the ntilum are presently under way. Institute. These workers suggested that in overall gross morphology (pinnule shape, size, ven-

ation, etc. ) the Naturita fossils are similar to FamUy POLYPODIACEAE fenis of the extant genus Bolbitis.

) reported fertile speci- Genus ASTRALOPTERIS Reveal, Tidwell Tidwell et al. ( 1967 and Rushforth mens of this fern and illustrated that this fern eould not be placed in the extant Bolbitis. Fur- Reveal, Astralopteris coloradica ( Brown ) thennore, since no living or fossil genus allowed Tidwell and Rushforth for the placement of this fern the new genus Astralojiteri.s- was proposed. Fig.s. 9-1, 9-7; 14-4; 19-4; 20. A. coloradica is one of the most common ele- 1950 Bohtis coloradica Brown-U.S. Geol. Surv. ments of the Westwater flora. In some collect- Prof. Paper 221-D, p. 49, PI. 12, Figs. 6-7. ing sites it is collected almost exclusively, al- 1967 Astralopteris coloradica ( Brovra ) Reveal, though it is most commonly collected in associ- Tidwell and Rushforth, Tidwell et o/.-BYU ation with Matonidium and Gleichenia. GoUec- Geol. Studies, V. 14, p. 2.39, PI. 2, Fig. 1; Pis. 3-6. tioas of fertile specimens of this fern are rare at 1968 Astralo])lcris coloradica ( Brown ) Reveal, the Westwater locality but, from recent exami- Tidwell and Rushforth, Rushforth and Tidwell- nations bv the author, appear to be somewhat Studies, v. 109, Pis. 1-3. BYU Geol. 15, p. more common from the Dakota Sandstone of Description.—Entire frond unknown; pinnae in- southwestern Golorado and Arizona. complete, large, ovate, tapering to rather acute apices; pinnules coriaceous, pinnate, alternate to opposite on rather stout rachises; lower pinnules UNGLASSIFIED FERNS linear-lanceolate, about 7 to 11 times as long as Brongniart broad, sessile to short stalked attaclunent; mar- Genus GLADOPHLEBIS gins entire; apices acute to rounded; upper pin- This generic n.mie was originally proposed lower ones, 4 to 5 times as long nules similar to by Brongniart (1849) but Saporta (1873) first as broad, basal attachment with rounded sinuses defined (Cladophlebis) as follows: in the uppennost pinnules; midvein prominent Frond pinnatelv cli\'idecl; pinnules separate to the pinnule apex or nearly so, arising decur- from one another, or slightlv miited, attached rently from rachis; lateral veins numerous aris- In the entire base. ing acutely and then becoming nearly at right angles to the midvein (or divaricate) for nearly Schimper (1874) emended this diagnosis to: their entire length, simple for one-half to three- Fronds pinnately divided; pinnae spreading; quarters or more of their length then usually bi- lolies or pinnules attached by the entire liase, furcating two or three times (or rarely trifur- sometimes confluent, rarely slightly auriculate, accuniinate or obtuse. occa.sional!)' dentate, cating at the first fork); ultimate veins finer, especialK- at the ape.\, not rarely subfalcately anastomosing, often obscure; sori round, biseri- curved upwards; midner\es pretty strong; sec- ate between the main lateral veins, about one- ondary nerves departing at a more or less fourth of the distance from the midrib to the acute angle, dichotomous; slender to very margin, fed from a costal areole and other sec- slender. ondary veins which extend to the placenta; spor- Fontaine ( 1889) modified this diagnosis only in angia 12 to 15, arranged in a ring around a cen- ". regard to venation by saying, . . midnerve tral placenta; annuli vertical; spores unknown. strong at base, and towards the summit dissolv- OccunRENCF..— Known from the Dakota Sandstone ing into branches."

) CAad- Formation near Westwater, Grand C;ounty, Fontaine ( 1889 discussed the usage of Utah. ophlebifi and concluded that it is a useful genus for placement of sterile foliage which may be Rei'Ositohy.— Brigham Young Universitv, BYU described as above. This is indeed the case, as 1415-1422; 1837-1840. foliage assigned to Cladopldebis could fit well

Discussion.—A. coloradica was originally de- into ;i luimber of different genera. Therefore, scribed as Bolhiti.s coloradica by Brown (19.50). Cladophlebis has become a useful, though un-

This placement was based upon sterile speci- niitunil genus which is very characteristic of the mens collected from Naturita, Montrose Gounty, Mesozoie, particularly of Jurassic times. Thus Golorado. Brown encountered difficulty in plac- Berrv ( 1912) mentioned that Cladophlebis is ing this fern and consefjuently he conferred essentially a fomi genus and includes species Biou)ck;al Skhies, \'ol. 14, No. 3 Dakoi a Sandstone Flora 35 ^^f^"^^ J

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Fig. 18. 1. MatonidiiiTii (?) hmcipinnulum Rusliforth. I^iriiiu fragment illustratini; pinnule dispositidn. (X1.2). Paratype: BYU 1570. 2. Matoniditim hrvunii Rusliforth. Trilete spore illii.strating weakly defined margo. (X1200). Paratvpe: BVU 1.561. 3. Mutonidium amcricanum Berry em. Rusliforth. Two pinna fragments. (X1.2). BVU 1.572. 4. Mutonidium aTiiericaninn Berry em. Rushforth. Portions of two pinnae illustrating venation. (X.3.6). BVU 1572. University Science Bulletin 36 Bricham Young

University, BYU which when fertile may be placed into several Hiii'osiTOHY.— Brigham Young different genera and even families. 1841. Fontaine (18S9) treated 23 species and sev- Discussion.—This feni is known from the West- varieties of Cladoplilchis from the American eral water flora only from a single specimen. This Group. However, Berry (1912) revised Potomac sp(>cimen illustrates fragments of two pinnules this genus, as known from the Potomac, and rec- each of which is rather well preserved, exhibit- eight species. As mentioned by Berry, ognizezd ing venation well. Fontaine recorded thi.s high number of species: This specimen agrees well with some speci- and . losing .sight of variation . . altogether mens placed by Fontaine (1889) in Cladophle- age or to position of the fos- changes due to bis constricta, especially PI. 2. Fig. 11a. How- sils with regard to the frond as a whole, as ever, due to the paucity of fossil material from well as changes due to the direct action of if the Utah speci- the environment. Westwater, it is not known mens agree in all details with the specimens Seward (1894) in treating Wealden fossils from Virginia. Even though this is the case, the Britich Museum of Natural History ac- in venation, overall size, and pinnule lobing, are all cepted the generic diagnosis of Schimper ( 1874) in agreement with C. constricta Fontaine em. Fontaine (1889). Berry (1912) as emended by Berry, and the author believes these similarities followed this same generic diagnosis, and also are sufficient to warrant the placement of the workers subsequent to Berry have done most Westwater fern in this species. likewise. The literature dealing with Cladophlethis is voluminous and is in need of reviewing. Many Cladophlebis parta Fontaine em. Berry species are recognized, although this is to be Figs. 8-5; 11-2. expected in anv unnatural genus as reproductive morphology may not be compared. However, 1889 Cladophlebis parva Fontaine-U.S. Geol. many species appear to be identical, and the Surv. Mon. 15, p. 73, PI. 4, Fig. 7; PI. 6, Figs. 1-3. genus should be monographed. 1899 Cladophlebis parva Fontaine, in Ward— U.S. Geol. Surv. 19th Ann. Rept., pt. 2, p. 657, Cladophlebi.s cotutricta Fontaine em. Berry PI. 160, Fig. IS.

Figs. 9-5; 11- 1. 1904 Cladophlebis parva Fontaine, in Ward— U.S. Geol. Surv. Mon. 48, 125, Pl. 65. Figs. 1889 Cladophlebis constricta Fontaine— U.S. p. 5-8. Geol. Surv. Mon. 15, p. 68, PI. 2. Fig. 11; PI. 3, Fig. 2; PI. 6, Figs. 5, 6, 8-14; PI. 21,>igs. 9, 13; 1911 Cladophlebis ])arva Fontaine, Berry— Mary- PI. 169, Fig. 2. land Geol. Surv., Low. Cret., p. 250, PI. 27, Figs. 1-2; Pis. 39-31. 1888 Chidophlehis latifolia Fontaine-U.S. Geol.

Surv. Mon. 15, p. 68, PI. .3, Fig. 1; PI. 6, Fig. 4. I'dl'lCladophlebls parva Fontaine, Berry— Proc. U.S. Nat. Mus., V. 41, 316. 1904 Cladophlebis constricta Fontaine, in Ward p. -U.S. Geol. Surv. Mon. 48, p. 197, PI. 71, Fig. 26. 1936 Cladophlebis ]>arva Fontaine, Bell—Geol.

Curv. Can. Mem. 285, p. 56, PI. 11, Figs. 3-5; PI. 1905 Cladophlebis virginensl's Fontaine, pro 12, Fig. 3; PI. 13, Figs. 1-2; PI. 14, Fig. 3; PI. 14, parte, in VVard-U.S. Geol. Surv. Mon. 48, p. 512, Fig. 3": PI. Ill, Fig. 7. Description.— Entire leaf unknown; pinnae prob- 91 Cladophlebis constricta l-'ontaine. Berry— ably lanceolate; rachis about 1 mm in diameter; Proc. U.S. Nat. Mus., v. 41, p. 314. pinnules alternate, linear-lanceolate, up to 26 Desciuition'.— Entire leaf unknown; pinnules mm long bv 6 mm wide, lobed approximateh' typically greater than 27 mm long by 10 mm one-half of the way to the midrib, margins of wide, margins deeply lobed up to one-half of the lobes sliglitlv undulate, apices acute, attach- distance to the rnidvein, lobe margins entire, ment basal to somewhat constricted basal; mid- lobe apices obtuse; midvein very strong; second- vein prominent, undi\ided to pinnule ;ipex; sec- ary veins repeatedly forked, open to the margin; ondary veins divide from one to thn-e times, re- fertile specimens unknown. main free; fertile specimens unknown.

Occurrence.—Near Westwater, Grand (^ountv, Occurrence. —Near Westwater, Grand County, Utah. Utah. a

Bi()L(k:ic:al Series, Vol. 14. No. 3 D.akot.\ S.^niwtone Fi.oh 37

Fig. 19. 1. Matonidium brounii Rushfortli. Pinna fninrriL-nt illustrating soriation and venation. (X3). Paratype: BYU 1563. 2. Malonidium (?) laiuijnniitilum Riishforth. Pinna fragment illii.strating pinnule disposition and venation. Holotvpe; BYU 1.569. 3. Matonidium hroivnii Rushforth. Pinnule exhibiting venation. (.\3). Paratype: BYU 1559. 4. Astralopteris cnlnradertsis Reveal, Tidwell, and Rushforth. Pinnule e.vhihiting venation. (X3). BYU 1837. 5. Matonidium americanum Berry em. Rushforth. Pinna fragment e.vhihiting venation. (X3). BYU 1831. 6. Matonidium brownii Rushforth. Pinna apex illustrating pinnule disposition. (X3). BYU 183.5. )

38 Brigham Young Univebsity Sciencf Bulletin

Repository.—Brigham Young University, BYU Family MAGNOLIACEAE 1842. Genus .MAGNOLIA L. Discussion.—Cladophlehi.s parva Fontaine em. Magnolia houlaijarui Lescjuereux

Berry is known only from one specimen in the FiR. 16-1. VVestwater flora. This specimen illustrates several pinnules on a pinna Iragment, a few of 1792 Maglunia hotddt/anu Lesquereux— U.S. which are rather complete. This fern from the Geol .Surv. Mon. 17. p.' 202, PI. 60. Fig. 2. VVestsvater locality has been assigned to Clcid- 1894 Ma

ANTHOPHYTA ( MAGNOLIOPHYTA Bull. Torrey Bot. Club, v. 36, p. 254.

Family AQUIFOLIACEAE 1911 Magnolia boulatjana Lesquereux, Berry-

New Jersey Geol. Surv. Bull. 3, p. 131, PI. 14, Genus ILEX L. Fig. 2.

Uex serrate Rushforth, sp. nov. Description.—Leaf greater than 7.5 cm long by

Figs. 12-5; 16-2. 4 cm wide, narrowly elliptic in outline, coriaceous; base rounded to bluntly rounded; margins Description.—Leaves vary from fairly short and entire; midvein strong, bearing somewhat in- wide long L7 wide) to longer (3.5 cm by cm fre(juent secondaries; secondary veias pinnate, and thinner cm long 1 wide), some- (4 by cm parallel, progressing to near margin and then what coriaceous, angular-ovate; apex acute to beinding to unite with vein above to fomi accuminate; base narrowing to the petiole; mar- rounded areoles near the margin; tertiary veins gins serrate to somewhat spinose; petiole short; connecting the secondaries. midvein weak, continuing to apex; secondary veins mostly obscured, borne at acute angles, Occurrence.—Near Westwater, Grand County, progressing to margin. Utah. Repository.- Brigham Universitv, Occurrence.—Dakota Sandstone Formation near Young BYU Westwater, Grand County, Utah. 184,3. Discussion.— Berry (1911) noted that leaves of Repository.—Brigham Young University, holo- Magnolia boulai/ana Lescjuereux are remarkably type: BYU 1801; paratype BYU 1892. constant in size and shape. This is particularly

Dlscu.ssion.—Leaves assigned to this species noteworthv when it is recalled that this con- from near Westwater are somewhat similar to stancy extends over wide geographical areas and Ilex dakotensis Les(juereux in shape and size. for a good deal of time. However, the leaf originally placed in l.dako- Berry (1911) noted that leaves assigned to tetm.'i dtx-s not exhibit serrate margins, and ven- Magnolia glaueoides Holliek by Holliek and ation differs somewhat from Utah specimens. In Newberry are identical to leaves of M. houlaij- overall shape, the Westwater material agrees ana. In view of this. Berry placed M. glaueoides rather closely with Ilex stenoplit/lla Unger from in synonomy under M. houtihii/ana. the Tertiary of Europe, as does Ilex dakotett^is. Placement of the Utah Magnolia in Magnolia However, Les(|uereux (1892) pointed out that houllayana Lesquereux may be made without these two species differ in size and somewhat in hesitation, as size, shape, and venation are all shape. In addition, /. stenoplujlla is not serrate identical between the two. to spinose as is the Utah Ilex. Ilex serrata differs from other species of Ilex Family MORACEAE in its small size coupled with spinose margins. Genus FICUS L. In margination, /. serrata is similar to /. armata Lesquereux, although the latter is much larger This genus in regard to the Mesozoic has be- and of different shape than /. serrata. come more of a form genus than a natural Biological Series, Vol. 14, No. 3 D.^ikota Sandstone Flora 39

^4»^'J3

' '.A?i^

Fic. 20. Astralopterui coloradica Reveal, Tidwell. and Rushforth. Pinna exhibiting pinnule disposition. (X1.2). BVU 1415. 40 Brigham Young University Science Bulletin

satisfaction, as might l)e expected for entire unit. Into this genus are placed leaves which leaves, the internal senation strucure of which are somewhat similar in that they exhibit laii- tends in m;iny towanl similarity. 1 regartl this However, Cretaceous raceous characteristics. reference to Fici/.v its makeshift. Many similar leaves placed in this gcmis often could be placed leaves from the Dakota Sandstone have been with ecjual logic in various other genera such as referred with little evidence to Andromeda. All are perhaps examples of lauraceous species Salix or Laurus. that are difficult to separate.

Ficus daplaiugenoidcs (Heer) Berry Family MYRTACEAE Figs. 16-3, 16-4. Genus EUCALYPTUS L. Herit 1867 Proteoide.1 daphnogenoides Heer, in Ca- Eucahjptus dakoten.'iis Les(juereux pellini and Heer—Denksch. Allgem. Schweiz. Fig. 1.5-8. Ges. Naturwiss., v. 22, p. 17, PI. 4, Figs. 9-10. 1892 Eucahjptus dakotensis Lesquereux—U.S. 1874 Proteoides daphnogenoides Heer, Lesquer- Geol. Surv. Mon. 17, p. 1.37, PI. .37, Figs. 14-19. eux— U.S. Geo!. Surv. Terr. Kept., v. 6, p. 85, PI. 15, Figs. 1-2. 1895 Eucahfptus angustifolia Newberry— U.S. Geol. Surv. Mon. 26, p. Ill, PI. .32, Figs. 1, 6, 7. 1905 Ficu.i daphnogenoides (Heer) Berry- Bull. Torrey Bot. Club. v. 32, p.329, PI. 21. 1895 Eucahjptus (?) nervosa Newberry— U.S. Geol. Surv. Mon. 26, p. Ill, PI. .32. Figs. .3-5, 8. 1921 Ficus daphnogenoides (Heer) Berry— U.S. Descriition.—Leaf linear, 4.5 cm long by 1.1 cm Geol. Surv. Prof. Paper 129, p. 163, PI. 39, Fig. 1. wide, somewhat coriaceous; base cuneate; mar- 1959 Ficus daphnogenoides (Heer) Berry, gins entire; midvein strong; secondary veins pin- BrowTi-U.S. Geol. Surv. Prof. Paper221-D, p. nate, arising obtusely, extending to leaf margin. 50, PI. 11, Figs. 1, 2, 4. Occurrence.—Near Westwater, Grand County, Description.—Leaves typically greater than 12 Utah. cm long bv 3 cm wide, more or less oblanceo- Repository.— Brigham Young University, BYU late; apex accuminate, fonning a rather long 1848. drip pohit; base cuneate; margins entire; mid-

vein strong at point of origin, becoming weaker Discussion.— Lesquereux ( 1892) described Eu- towards leaf apex; secondary veins pinnate, a- cahjptus dakotensis as follows: rising acutely, bending to parallel margin; terti- Leaves coriaceous, linear, or gradually narrow- ary veins obscured. ed from an obtuse apex to the base, deeur- Occurrence.—Near Westwater, Grand County, ring into short, alate petiole; borders recur\ed, medi;m nerve strong; secondaries thin, obli(|ue. Utah. proximate, parrallel. camptodrome. Repository.- Brigham Young University, BYU Lesquereux further pointed out that the margin 1844-7846. of some specimens was flat rather than recurved. Discussion.—Lesiiuereiix (1874) described Pro- Eucahjptus angustifolia Newberry and £. (?) teoides daphnogenoides Heer (Ficus daphno- nervosa Newberry appear to be specifically genoides) as follows: identical both with each other and with E. dakotensis Lescjuereux. The only apparent difference Lfiivcs oviitc-laiiccolatf near the base, j^radii- is E. is naked ally tapering uiiuard to a long, aeiite, seythe- that the petiole of angustifolia shaped point, entire, smooth, and coriaceous; rather than a];ite. medial ner\e narrow; secondary veins obsolete, The Westwater Eucahjptus also agree well few, ascending under a verv acute angle from with the description of Les(|uereux (1892), ex- the medial nerse and following the borders. cept that the petiole of the Utah sjjecimen is This description fits the Westwater Ficus in all not winged. However, this difference does not respects, and the identity of the two may bo warrant specific separation, and E. dakotensis

fairly certain. is used for placement of the Westwater speci-

Berry ( 1905) transferred Proteoides daphno- men.

genoides Heer to the genus Ficus. This trans- As mentioned by Les((uereux ( 1892) E. dako- ferral was valid, ;dtliongh Brown (1950) re- tensis differs from an ;ipp;ireiitlv closely rei;ited

marked: fonii, £. geinitzi in that the tonner is snuiller

These entire-margined leaves li.ivc been re- and linear in shape as opposed to larger and ferred to manv different genera with little broadly spatulate in shape. Biological Sehies, \'ol. 14, No. 3 Dakota Sands ione Flora 41

t ^ 1* * a X 'W:-*^!

x s } "V -.,^ ^*^v

; .^.i* «??v a

•^.'

*?, ^ 'V-' <^' '^^> \Ul»'

" 4 ;^, ^^.

'^, ^A,^ w ^,

Fic. 21. A. Gk'ichenia conipfoniticfolia (Deb. and Ett.) Heer. Pinna fragments e.xhibiting pinnule dispo.sition. (XI. 5). BVU 1853a. B. Mutoiiidium hroicnii var. magnipinnuluni Hiishforth. Pinna fragments. (XI. 5). BYU 18.53b. Science Bulletin 42 Bbigham Young University

Family PLATANACEAE Occurrence.—Dakota Sandstone Formation near Rabbit Valley, Grand County, Utah. Genus PLATANUS L. Rkpository.— Brigham Young University, BYU the Cre- Leaves of the genus Platanus from 1849. taceous are well known and rather common. Discussion.—This leaf agrees in all details with Manv of these leaves are essentially indistin- Plantanus neuherryana Heer, and is very similar guishable from those of extant species (Seward, to leaves figured under this species by Lesquer- 1827). eux and Seward. In addition, it is very similar to leaves described as Platanus affinis, Pro- Platanus newbernjana Heer tophijllum minus, and ProtophtjUum nebrascense

Fif;. 16-6. by Lesquereux (1874, 1882). The Westwater leaf exhibits a rounded to angular base, which 1867 Platanus newbernjana Heer, in Capellini agrees with the description of Platanus new- and Heer—Denksch. Allgem. Schweiz. Ges. Na- bernjana by Lescjuereux ( 1874 ) , although in turwiss., V. 22, p. 16, PI. 1, Fig. 4. this respect it differs somewhat from specimens 1868 Platanus neioberrtjana Heer. Lesquereux— placed in this species by Seward ( 1927 ) which Amer. Jour. Sci., v. 46, p. 97. exhibited very rounded bases. 1874 Platanus netLhern/aiui Heer, Lesquereux— Seward (1927) noted that leaves of Platanus

U.S. Geo!. Surv. Terr. Rept., v. 6, p. 72, PI. 8, newbernjana "agree very closely with the recent Figs. 2-3; PI. 9, Fig. 3. Platanus mexicana Moric."

1874 Platanus affinls Lesquereux— U.S. Geol. Surv. Terr. Rept., v. 6, p. 71, PI. 4, Fig. 4. FamUy SALICACEAE

1873 ProtophtjUum minus Lesquereux— U.S. Genus SALIX L. Fig. 1. Geol. Surv. Terr. Rept., v. 6, p. 194, PI. 27 Salix newbernjana Hollick 1874 Protojiht/lluin ncbrascense Lesquereux— Figs. 12-4; 15-7. U.S. Geol. Surv. Terr. Rept., v. 6, p. 103, Pi. 27, Fig. 3. 1895 Salix newbernjana Hollick, in Newberry— U.S. Geol. Surv. Mon. 26, p. 68, PI. 14, Figs. 1-7. 1882 Platanus affinis Lesquereux, Heer— Flor. 1911 Salix newbernjana Hollick, Berry—New Foss. Arct., V. 6, pt. 2, p. 73, PI. 18, Figs. 16-17. Jersey Geol. Surv. Bull. 3, p. 113, PI. 11,' Fig. 2. 1883 Platanus newbern/ana Heer, Lesquereux— Description.—Leaves greater than 6 cm long by U.S. Geol. Surv. Terr. Rept., v. 8, p. 28, PI. 59, up to 2.5 cm wide, lanceolate; base rounded; Figs. 1-6; PI. 60, Fig. 1. apex accuminate to fonn a rather well defined 1883 Cissites affinis Lesquereux— U.S. Geol. drip point; margins finely crenate to dentate; Surv. Terr. Rept., v. 8, p. 67. midvein fairly strong; other veins obscure. 1927 Platanus neiiberrtjana Heer, Seward— Phil. Occurrence.—Dakota Sandstone Fonnation near Trans. Rov. Soc-. London B, v. PI. 215, p. 128, Westwater, Grand County, Utah. 11, Fig. 116; Text- Fig. ,30. Repository.— Brigham Young University, BYU Description.—Leaf greater than 5 cm long by 3 1850-1851. cm wide, coriaceous; base rounded; midvein strong bearing numerous secondary veins at Discussion.—This leaf from Westwater is essen- acute angles; secondary veins apparently extend tially identical to those described by Hollick (in to ( margin caspcd(xlrome ) ; tertiary veins nu- Newberry, 1895) and Berry (1911). It may be merous, borne at nearly 90° angles to secondary placed with confidence in Salix twwbernjana veins, connecting secondaries. Hollick.

ACKNOWLEDGMENTS

The author wishes to express his appreci- also due to several people who aided the author ation for the many helpful suggestions of Dr. in completing field work and technical aspects W. D. Tidwell and Dr. R. J, Bushman during of this manuscript. the preparation of this manuscript. Thanks are Biological Sehies, Vol. 14, No. 3 D.\kot.\ S.\ndstone F"lora 43

REFERENCES CITED

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. 1958. Heniarks on the Lower Cretaceous beds vironment of the angiosperms: Amer. [our. Bot.. v.

of Kansas and Nebraska, togetlicr with descriptions 2, no. 1, p. 1-22, 4 Pis. Carboniferous fossils of some new species of from Stanton, T. W. 1905. The Morrison Formation and the valley of the Kansas River: Proc. Phil. Acad. its Relations with the Comanche Series and the Da- Nat. Sci., V. 10, p. 256-260. kotii Formation: Jour. Geology, v, 13, p, 657-669, . 1859. On the so-called Triassic rocks of Kan- Stephenson, L. W, 1952, Larger invertebrate fossils sa.s and Nebraska: Amer. Jour. Sci. and Arts, v. 27, of the Woodbine Formation (Cenomanian) of p. 31-35. Texas: U.S, Geol, Surv. Prof. Paper 242, 226 p.. 1861. Descriptions of new Lower Silurian 51 PLs, (Primordial), Jurassic, Cretaceous, and Tertiary fossils collected in Nebraska Territorv. with some Stokes, W. L, 1948. (leologv of the Egnar-Gypsum remarks on the rocks from which they were obtain- V'allev area, San Miguel and Montrose Counties. LI.S. Surv. Invest. Pre- ed: Proc. Phil. Acad. Nat. Sci., v. 13, p. 417-432. Colorado: Geol. Oil and Gas S. 1868. lim. Map 93. Newbehrv, J. The late extinct floras of North .•Vmeric:!: .^nn. Neu York Lvceum Nat. Hist., v. 9, Tester, A. C. 1931. The Dakota Stage of the type p. 1-76, localitv: Iowa Geol. Surv. Ann. Rept. 35, p. 200-

. 1895, The flora of the Ambov Clavs: U,S, 332.

Geol, Surv. Mon. 26, 258 p. TnnvELL. S. R. L. Reveal. W, D., Rushforth and J. 1898. The later extinct floras of Nortli 1967. Astraloptcris, a new Cretaceous feni genus U.S. Geol. America: Surv. Mon. 35, 295 p. from Ut:ili :uul Colorado: Brigham Young Univ. PiEHCE, R. L. 1961. Lower Upper Cretaceous phuit Geol. StnBlack Mesa, Navajo and Velenovsky, 1888. Die der Bohinischen Hopi Indian Reservations, Arizona: Amer. Assoc. |. fame Kreitlformation: .Abh. K. Bohm. Ges. Wiss., Math- Petroleum Gm\. Bull. \. 40, No. 2, p. 2.55-294. Natw., ser. 7. v. 1-.32, 6 Pis. RicHTEH, p. B. 1906. Beitrage zur flora der Unteni 2. p.

Kreide (^uedlinburgs, I. Die gattnng Hau.smuitnid W'ahi), L. F. 1899. The Cretaceous formation of the

Dunker unci eiuige seitenere pflanzenreste: 25 p., Black Hills as indicated bv the fossil plants: U.S. 7 pLs., Leipzig, Geol. Sur\. 19th Ann, Rept,, pt. 2. p. .527-.551, RusHFORTii, S. R, and W. D. Tidewell. 1968. Notes 702-810.

on the distribution and morphology of the fern . 1995. Status 111 the Mesozoic Floras of the

genus AstruloptiTis: Brigham ^'oung Uni\. Cleoi. United States; U.S. G.'ol. Sur\ . Mon. 48, 616 p„ Stud., V. 15, pt. 1, p. 1(W-115. 119 Pis.