Taxonomic Revision of the Spermatopsida 

of the Oligocene Creede Flora, 

Southern Colorado 

U.S. GEOLOGICAL SURVEY BULLETIN 1923 
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Taxonomic Revision of the Spermatopsida 

of the Oligocene Creede Flora, 

Southern Colorado 

By JACK A. WOLFE and HOWARD E. SCHORN 

U.S. GEOLOGICAL SURVEY BULLETIN 1923 
DEPARTMENT OF THE INTERIOR 

MANUEL LUJAN, JR., Secretary 

U.S. GEOLOGICAL SURVEY 

Dallas L. Peck, Director 

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purposes only and does not imply endorsement by the U.S. Government. 

UNITED STATES GOVERNMENT PRINTING OFFICE: 1990 

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Second printing 1991 

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Library of Congress Cataloging-in-Publication Data 

Wolfe, Jack A., 1936-

Taxonomic revision of the Spermatopsida of the Oligocene Creede flora, 

southern Colorado I by Jack A. Wolfe and Howard E. Schorn. 

p. em. - (U.S. Geological Survey bulletin; 1923) 

Includes bibliographical references. 

Supt. of Docs. no.: I 19.3:1923 

1. Paleobotany-Oligocene. 2. Paleobotany-Colorado. 3. Creede 

Formation (Colo.) I. Schorn, Howard E. II. Title. Ill. Title: Spermatopsida of 

the Oligocene Creede flora, southern Colorado. IV. Series. 

QE75.89 no. 1923 

[QE928] 

557.3 s-dc20 9G-3265 

[561 I .19788] CIP 
CONTENTS 

Abstract 1 

Introduction 1 

Systematics of the Creede flora 3 

Systematic list 4 

Systematic descriptions and discussions 4 

Cupressaceae 4 

Pinaceae 5 

Berberidaceae 15 

Salicaceae 18 

Philadelphaceae 19 

Grossulariaceae 20 

Rosaceae 23 

Leguminosae 32 

Bignoniaceae 33 

Liliatae 34 

Names of dubious value 34 

Summary of systematic revisions 36 

Description and discussion of a Neogene spruce 37 

References cited 38 

PLATES 

[Plates follow text] 

1. Juniperus, Abies, and Pinus. 

2. Pinus and Pinaceae indetermined. 

3. Pinus and Picea. 

4. Berberis and Mahonia. 

5. Populus, Jamesia, and Ribes. 

6. Ribes. 

7. Eleiosina. 

8. Eleopoldia. 

9. Ribes, Holodiscus, Sorbus, and Crataegus. 

10. Cercocarpus and Potentilla. 

11. Potentilla, Prunus, Rosoideae gen. indet., and Osmaronia? 

12. Catalpa, Prunus, and indeterminate leaves. 

13. Pinus, Eleopoldia, and achenes of uncertain affinity. 

FIGURES 

1. Diagram showing proposed terminology for winged seeds of Pinaceae 7 

2-3. Illustrated keys for generic identification of winged seeds of Pinaceae that 

have: 

2. Adfacial surface exposed 7 

3. Abfacial surface exposed 8 

Contents Ill 
4-5. Histograms of: 

4. Needle length for five-needled fascicles of Creede Pinus 9 

5. Leaf length in Creede Abies 10 

6-9. Drawings showing: 

6. General terminology of apophysis, umbo, and mucro of ovulate cone 

scale in Pinus 12 

7. Ovulate cones of Creede species of Pinus 13 

8. Areolar venation of Mahonia 17 

9. Venation of Populus larsenii 18 

10. Histogram of laminar dissection in Ribes lacustroides 21 

11-13. Drawings showing: 

11. Venation of Osmaronia 31 

12. Venation of Prunus 32 

13. Basal configuration of Cercis 32 

TABLES 

1. Abrasion stages of Creede ovulate cones of Pinaceae 6 

2. Differentiation of Creede species of Ribes 20 

IV Contents 
Taxonomic Revision of the Spermatopsida of the 

Oligocene Creede Flora, Southern Colorado 

By Jack A. Wolfe and Howard E. Schorn1 

Abstract 

Determination of impressions and compressions of the 

Oligocene Creede flora of Southern Colorado by two different 

methods results in markedly different taxonomic results. One 

method, based on ·matching" extant plant organs with 

fossils, gives little or no consideration to taxonomic 

significance of the ·matching" characters in extant plants. 

Many ·matches," especially of poorly preserved material are 

based on the present-day association of the •matched" taxon 

associated with taxa represented in the fossil assemblage by 

well-preserved material. Some •matching• characters have 

resulted from different abrasion and (or) degradation stages, 

and other "matching• characters represent different stages in 

ontogenetic development. The second method considers all 

these significant factors, and application of the method to the 

Creede fossils results in major reduction in numbers of 

families, genera, and species, as well as reassignment of 

many fossils to different supraspecific categories. 

INTRODUCTION 

The Creede flora represents the only documented 

late Oligocene plant assemblage from the southern 

Rocky Mountains. Hence, valid determinations of the 

Creede taxa are critical to understanding biogeographic 

patterns, phylogeny of clades represented in the Creede 

flora, and the paleoecologic implications of the Creede 

assemblage (Wolfe and Schorn, 1989). The geologic 

setting of the Creede plant -bearing beds, which were 

deposited in the moat of a calderon, has been discussed 

Manuscript approved for publication, December 13, 1989. 

1

Museum of Paleontology, University of California, Berkeley, 

California. The authors contributed equally to this report. Authorship 

is not alphabetical because Geological Sutvey policy requires that the 

first author be a Sutvey employee, contractor, grantee, or volunteer. 

elsewhere (Axelrod, 1987; Wolfe and Schorn, 1989). The 

Creede plant -bearing beds are radiometrically dated at 

27.2 million years (Steven and others, 1967; corrected for 

newer decay constants). 

A recent taxonomic treatment of the Creede flora 

recognized 73 species of Spermatopsida (Axelrod, 1987). 

Recognized were 2 families, 6 genera, and 19 species of 

conifers; we recognize 2 families, 4 genera, and 8 species. 

Also recognized were 21 families, 37 genera, and 53 

species of dicotyledons; we recognize 7 families, 18 

genera, and 24 species. Many of Axelrod's (1987) 

species, moreover, are reassigned to different families 

and (or) genera. How can two systematic treatments of 

the same material result in such markedly different 

systematic dispositions of so many taxa? 

Axelrod's (1987) systematic treatment of the 

Creede flora is fundamentally "picture matching" 

(Wolfe, 1974). In this method, for a given fossil leaf, an 

extensive search is made of modern comparative 

collections in herbaria to find the closest "match" for the 

fossil. The fossil is then assigned to the same genus, 

family, and order as the modern "match"; these 

assignments are not made because the fossil has 

characters that circumscribe the genus, family, or order. 

The characters that result in the "match" are typically not 

stated, and, even if they are, no evaluation is made as to 

whether the characters are taxonomically critical (as 

opposed to characters that might represent ontogenetic 

response to environment). If foliar characacters (or 

characters of other organs) are described and compared, 

the descriptions and comparisons are typically phrased in 

undefined terms or terms of uncertain meanings. 

Examples of "matches" that are based on uncritical 

comparisons abound in literature on western American 

Tertiary floras. Axelrod (1964, p. 127) described samaras 

assigned to Acer bolanderi Lesq. (sect. Acer) as having 

"***a wing that is sharply truncated under the nutlet, and 

the latter displays a complex network of interlacing 

prominent veins" versus samaras assigned to A. columbianum Chan. & Axelr. (supposedly sect. Glabra) as 

Introduction 
"***having a deep sinus under the nutlet owing to 

constriction of the wing there, and the nutlet does not 

display a network of veins but has a deep prominent 

wrinkle." If these fossil samaras are so clearly 

distinguishable and each is "matched" by different 

species in different sections of Acer, then how was it 

possible for one specimen (Axelrod, 1964, pl. 15, fig. 9) 

to be assigned to A. bolanderi and the counterpart 

(Axelrod, 1964, pl.16, fig.10) ofthe same specimen to be 

assigned to A. columbianum? 

A method that is frequently employed to narrow 

the search for a "match" is based on modern associations. 

Once a few to several taxa (typically the most abundant) 

are determined in a fossil flora, if the presumed 

"matches" of these taxa now live in proximity, then search 

is made among the presently associated taxa, or at least 

those living in the same general area. Poorly preserved 

fossil leaves are especially prone to be "identified" in this 

manner; the "identifications" are difficult, if not 

impossible, to falsify, but they are seemingly consistent 

with the modern association. Coming full circle, such 

"identifications" have been used as evidence for associational stability during the Cenozoic. A classic example 

of how associational "matches" can deny morphologic 

data is that of a taxon originally described as Cinnamomum bendirei in the Oligocene Bridge Creek flora of 

Oregon. As discussed by Wolfe (1969, p. 86): 

The ·holotype• of the associational method of determination (Cain, 1944, p. 43) is in fact an excellent demonstration of the weakness of the method. Knowlton 

(1902) originally described Cinnamomum bendirei 

from the Bridge Creek assemblage of Oregon. Chaney 

(1927), however, interpreted this assemblage as a redwood forest and thus considered Cinnamomum an incongruous element. He therefore transferred Knowlton's species to Philadelphus, which would be expected in a redwood forest. Brown (1940) pointed out 

that the leaves morphologically could not be Philadelphus but had the diagnostic characters of Lauraceae; 

he thus transferred the species to Sassafras. This 

transfer was ignored by Chaney (1944b, p. 350), who 

still considered that Philadelphus was a better choice 

for a redwood forest. After the discovery of the living 

Metasequoia and the realization that the Bridge Creek 

assemblage contained not Sequoia but rather Metasequoia, Chaney (1952) accepted Brown's transfer of the 

species to Sassafras. The leaves, however, have the 

small areolas lacking branched, freely ending veinlets 

and the continuous marginal [now fimbria!] vein of 

Cinnamomum, Lindera, and Neolitsea; Sassafras has 

large areolas intruded by branching veinlets and lacks 

a marginal vein (Wolfe, 1960). Knowlton's determination was more valid than either Chaney's or Brown's. 

Both the "picture matching" and "associational" approaches to determinations of fossil leaves, especially 

those of the dicotyledons, have led to many similar misidentifications. 

2 Spermatopsida of Oligocene Creede Flora, Colorado 

A second source of error in misidentifications of 

fossil dicotyledonous leaves is misapplication of the 

"living equivalent" concept. The worker basically 

considers the extant species that has foliage (or other 

organ) most similar to the fossil as the "living equivalent" 

of the fossil taxon. This concept was originally developed 

for purposes of inferring paleoecology and paleoclimate 

by assuming that morphologic similarity indicated 

physiologic (and hence ecologic) similarity. We will not 

here discuss ecologic problems inherent in this 

assumption but will emphasize the taxonomic problems 

that misapplication of the "living equivalent" concept has 

caused. 

The misapplication is assuming that all fossil leaves 

(or other organs) that have the same supposed "living 

equivalent" represent the same fossil species. MacGinitie 

(1953, p. 79) pointed out the obvious: two fossils can both 

be closest morphologically to the same living species and 

yet can differ morphologically from that living species in 

very different directions. For example, Wolfe (1964) 

assigned certain Acer samaras from the Miocene of 

Nevada to the extant A. macrophyllum Pursh (the sole 

extant member of section Macrophylla) and synonymized 

with this species the fossil species A. oregonianum 

Knowlt. Detailed morphologic analysis, however, 

indicates that the Nevada samaras represent an extinct 

lineage very distinct from that of A. oregonianum, which, 

in turn, represents an extinct lineage distinct from that of 

its supposed "living equivalent" A. macrophyllum, 

although all three lineages belong to Macrophylla (Wolfe 

and Tanai, 1987). 

A second point relevant to the "living equivalent" 

concept is: a fossil species can have an uncanalized 

morphology (Upchurch, 1985); that is, a fossil species 

can have wide morphologic variation, even wider than the 

variation possessed by extant taxa. If this variable fossil 

taxon gave rise to several extant taxa by canalizing 

different characters in different directions, then a 

number of subsets of the fossil taxon will be most similar 

to a number of the descendant taxa; misapplication of the 

"living equivalent" concept will then lead to the 

partitioning of the fossil taxon into a number of fossil 

species (see later discussion of Ribes lacustroides). 

Excessive partitioning of a fossil taxon at the 

generic level can also result from misapplication of the 

"living equivalent" concept. In the rosaceous Chamaebatiaria group, Wolfe and Wehr (1988) described a middle 

Eocene taxon, Stonebergia columbiana, from one locality 

in British Columbia. The variation of S. columbiana 

includes characters of Chamaebatiaria, Chamaebatia, 

and Sorbaria, and Stonebergia is considered to be 

ancestral to all three extant genera. Alternatively, all 

three extant genera could be considered as "living 

equivalents" of the most similar fossils. Specimens of 

Stonebergia columbiana that are most similar to Sorbaria, 
for example, might form the basis for a species of 

Sorbaria and the other fossil specimens assigned to two 

additional species, one placed in Chamaebatiaria and the 

other in Chamaebatia. As invalid as this alternative 

taxonomic partitioning might seem, the same logic has 

been applied to determinations of numerous Tertiary 

fossils. 

In misapplication of both the "associational" and 

"living equivalent" methodologies, the extant leaf (or 

other organ), rather than a fossil, has become the bolotype for the morphologic definition of a fossil species. 

Comparisons then may not be made between two fossils 

but rather between each fossil and the extant organ, and, 

if both fossils are similar to the organ(s) of the same 

extant species, the two fossils are considered to be 

con specific. A major result of these approaches has been 

that morphologic differences between fossil and living 

taxa or between two fossil taxa are ignored or lost; this 

morphologic information may be critical in phylogenetic 

reconstructions and in understanding relationships 

between extant plants. 

For many decades these approaches were the primary methods used for identification of Cretaceous and 

Tertiary foliage. Since the 1960's and 1970's, however, 

paleobotanists working with leaves have developed (1) 

criteria for the recognition of characters that are critical 

at taxonomic levels above the species and (2) rigorous 

terminology for descriptions (for example, Hickey, 1973; 

Dilcher, 1974; Wolfe, 1974, 1989; Hickey and Wolfe, 

1975). Our taxonomic revision of the Creede flora 

reflects this newer approach to the determinations of 

fossil leaves. Development of this new approach, 

commonly referred to as leaf architecture, involves 

detailed comparative morphologic and (or) anatomical 

analysis of a wide range of foliage of extant plants, 

typically using chemically cleared specimens. 

Taxonomically critical foliar characters are those that 

circumscribe groups of taxa as traditionally recognized 

on the basis of criteria of morphology of reproductive 

structures. 

Acknowledgments.-We are indebted to Peter 

Robinson for access to the University of Colorado's 

Creede collections and to Scott Wing for access to the 

Smithsonian Institution's Creede collections. The part of 

the report dealing with conifers was critiqued by C. 

Millar and the late W.B. Critchfield (U.S. Forest Service) 

and that dealing with angiosperms by G.R. Upchurch 

(National Center for Atmospheric Research) and S.L. 

Wing (Smithsonian Institution). Harvey Scudder 

(California Academy of Sciences) offered considerable 

help regarding supposed plant structures that actually 

represent insect remains. We have also benefited by 

discussions with James Eckenwalder (University of Toronto) and Patrick Fields (Michigan State University). 

Baki Kalispagil (University of California, Berkeley) 

supplied the cone of Pinus from the Miocene of Turkey. 

Modern comparative material was supplied by the 

following herbaria: A, G, CAS, MO, UC, and US. 

SYSTEMATICS OF THE CREEDE FLORA 

Many of the Creede species described as new by 

Axelrod were minimally described, and thus we include 

thorough descriptions based (for the angiosperms) 

primarily on Hickey's (1973) terminology. Exceptions 

and (or) additions are: (1) "abmedial" is used in place of 

Hickey's "exmedial" (Wolfe and Wehr, 1987), (2) 

"external secondary vein" (von Ettingshausen, 1861) is 

used in preference to Hickey's "secondary vein branch," 

(3) topology of teeth and their associated veins is given 

(Hickey and Wolfe, 1975), (4) in lobed leaves, topology 

oflobal sinuses and their associated veins is given (Wolfe 

and Tanai, 1987), (5) veinlets that branch irregularly 

from a few to several times are referred to as "dendritically branching," and ( 6) descriptive terms for deeply 

cleft laminae follow Wolfe and Wehr (1988). 

Comparisons of various conifer organs would be 

greatly facilitated if all workers adopted a standardized 

approach for illustration. By convention, objects should 

be oriented with their upper/distal/apical ends toward the 

top of a page or plate. Similarly, a standardized 

terminology for each conifer organ would allow ready 

comparisons, even in the absence of illustrations. For 

some organs, terminology exists in the literature, but the 

winged seeds and cone scales of, for example, Pinus have 

received much less attention. Klaus (1980), however, 

deals thoroughly with apophysis-umbo-mucro relations 

for Pinus (see fig. 6 later) cone scales, and we propose 

(see fig. 1 later) a terminology for pinaceous winged 

seeds. 

If standardized morphologic terminologies are not 

employed and particularly if descriptions are very brief, 

we urge that illustrations of specimens be adequate for 

other workers to recognize critical characters of the 

taxon. In many instances, this should involve photographic enlargements and (or) line drawings at the 

appropriate magnification to illustrate critical characters. 

Some of Axelrod's (1987) new species were also 

improperly typified according to the International Code 

of Botanical Nomenclature (ICBN) (Greuter and others, 

1988). We have also suggested corrections for some 

orthographic errors in spelling of specific epithets 

according to the recommendations of the ICBN. 

We have generally excluded from the study most 

specimens of presumed angiospermous reproductive 

structures that were determined by Axelrod (1987); most 

of these specimens are poorly preserved impressions 

that lack characters critical for identification. The 

Systematics of the Creede Flora 3 
infructescence assigned to Populus (Axelrod, 1987, 

pl. 21, fig. 1), however, has sufficient characters 

preserved to indicate probable validity of the generic 

assignment. The specimen thought to represent a pod of 

the legume Robinia (Axelrod, 1987, pl. 32, fig. 6) also has 

critical characters preserved; this specimen represents a 

larva of a taxon of the crane-fly family Tipulidae (H.I. 

Scudder, oral commun., March 1989). 

As the last item in each synonymy, the designation 

used by Wolfe and Schorn (1989) is given. A few 

taxonomic changes were made after the Wolfe and 

Schorn (1989) manuscript was accepted for publication. 

These changes involve recognition of two additional but 

rare species of Pinus and assignment of most fascicles of 

Pinus toP. crossii. These changes only minimally alter the 

sample scores or scores of other species in the multivariate analyses. 

The specimens on which our revision of the Creede 

flora is based include the material deposited in the 

University of California Museum of Paleontology 

(UCMP) and the University of Colorado Museum 

(Geology Department) (UCMG); that is, the same 

material that formed the basis for Axelrod's (1987) 

treatment. Material, mostly collected by B.K. Stewart in 

the 1930's, housed in the U.S. National Museum 

(USNM) and material in the Harvard University 

Botanical Museum (HUBM) has also been used. 

Included in these collections is material of one taxa 

(Rosoideae gen. indet. ) that was apparently overlooked 

in the prior systematic analysis. 

Systematic List 

Coniferophyta 

Coniferae 

Coniferidae 

Coniferales 

Cupressaceae 

Juniperus creedensis Axelr. 

Pinaceae 

Abies rigida Knowlt. 

Picea sp. 

Pinus crossii Knowlt. 

Pinus riogrande Axelr. 

Pinus sanjuanensis Axelr. 

Pinus sp. 1 

Pinus sp. 2 

Magnoliophyta 

Magnoliatae 

Ranunculidae 

Ranunculanae 

Berberidales 

Berberidaceae 

Berberis coloradensis Axelr. 

Mahonia aceroides (Knowlt.) Schorn & Wolfe 

Dilleniidae 

Dillenianae 

4 Spermatopsida of Oligocene Creede Flora, Colorado 

Salicales 

Salicaceae 

Populus larsenii (Knowlt.) Schorn & Wolfe 

Salix sp. 

Rosidae 

Rosanae 

Saxifragales 

Philadelphaceae 

Jamesia caplanii Axelr. 

Grossulariaceae 

Ribes lacustroides Axelr. 

Ribes robinsonii Schorn & Wolfe 

Ribes obovatum Schorn & Wolfe 

Rosales 

Rosaceae 

Eleiosina praeconcinna (Cocker.) Schorn & Wolfe 

Eleopoldia lipmanii (Axelr.) Schorn & Wolfe 

Stockeya creedensis (R. W. Br.) Wolfe & 

Wehr 

Holodiscus stevenii Schorn & Wolfe 

Crataegus creedensis Axelr. 

Sorbus potentilloides (Knowlt.) Axelr. 

Potentilla creedensis (Axelr.) Schorn & Wolfe 

Cercocarpus henricksonii Schorn & Wolfe 

Cercocarpus nanophyllus Schorn & Wolfe 

Rosoideae, gen. & sp. indet. 

Osmaronia? stewartiae (Axelr.) Schorn & Wolfe 

Pntnus creedensis Axelr. 

Pntnus sp. 

Fa bales 

Leguminosae 

Cercis sp. 

Legume, gen. & sp. indet. 

Asteridae 

Lamianae 

Scrophulariales 

Bignoniaceae 

Catalpa coloradensis (Axelr.) Schorn & Wolfe 

Liliatae 

Incertae Sedis 

Monocotylophyllum sp. 

Systematic Descriptions and Discussions 

Family CUPRESSACEAE 

Genus JUNIPERUS Linnaeus 

Juniperus creedensis Axelrod 

Plate 1, figures 1, 2 

Junipems creedensis Axelrod, 1987, p. 99, pl. 16, figs. 1-9. 

Sabina linguaefolia auct. non (Lesquereux) Cockerell. Knowlton, 1923, p. 187. 

Wolfe and Schorn (1989) designation: Juniperns creedensis. 

Description. -Foliated axis poorly preserved, 

length about 9 em, width of basal axis 0.4 em and of apical 

parts about 0.15 em; branching alternately (apparently 

from all sides of axis) .. Leaf shape rhombic, ternate 

phyllotaxis common, decussate phyllotaxis rare and 

typically in more apical parts of axis, infrequent poorly 

expressed glands. Fruit globose, diameter 0.9-1.4 em, 

detailed preservation lacking but seed cavities present, 

stock 0.1-0.2 em long and with minute leaves. 
Discussion. -Juniperus creedensis was described 

(Axelrod, 1987, p. 99) as "***scale leaves in 2's [decussate], opposite, closely appressed * * * ." Most specimens 

of J. creedensis have ternate rather than decussate phyllotaxis. The poorly preserved holotype of J. creedensis 

shows rare glands and both ternate and decussate phyllotaxis. The ternate phyllotaxis is most prominent in 

older (proximal) portions of the axis and is particularly 

evident in specimens that preserve a natural break of the 

axis. The rare, more flattened decussate phyllotaxis 

occurs in younger (distal) portions of the foliage. The 

tern ate phyllotaxis excludes the Creede material from the 

New World group of Cupressus, which has exclusively 

decussate phyllotaxis. 

Juniperus creedensis was compared to the extant J. 

osteosperma (Torr.) Little and the extant J. califomica 

Carr. (Axelrod, 1987). Juniperus osteospenna is both 

decussate and ternate, and the glands are typically poorly 

expressed (Edwards, 1983), as in J. creedensis; J. californica is mostly ternate and has well-developed glands. The 

Neogene J. nevadensis Axelr. has phyllotaxis and degree 

of glandularity similar to J. creedensis, but the axes and 

ovulate cones of J. creedensis are slightly more robust 

than in J. nevadensis (Axelrod, 1987). 

A second described species of Creede Juniperus, J. 

gracillensis Axelr., is not a gymnosperm. Although 

described as having closely appressed, scalelike leaves 

that have two-ranked (decussate) phyllotaxis (Axelrod, 

1987, p. 100), the specimens are branching, planar laminae that have medially positioned vascular strands. These 

specimens represent the rosaceous Eleopoldia lipmanii 

(pl. 13, fig. 3). 

Holotype.-7447. 

Paratypes .-7448-7475. 

Family PINACEAE 

Discussion.-The extensive collections available 

from the Creede area (Axelrod, 1987, tables 1-4) 

produced 7,628 conifer specimens. Almost half (3,668) 

are impressions of foliage axes and cones of a single 

species of juniper. The remaining half (3,955) Axelrod 

distributed among 10 species of Pinus, 3 species of Picea, 

2 species ofAbies, and 1 species each ofPseudotsuga and 

Tsuga. 

The assignment of the Creede material to 5 genera 

and 17 species of Pinaceae exaggerates the diversity of 

the family at Creede and partially resulted from failure to 

exercise caution regarding the identification of poorly 

preserved material. Identification of the various 

impression organs must rely on adequate morphological 

information; that is, in order for any fossil to be assigned 

to a particular taxonomic level, the fossil must have the 

set of characters that are necessary to identify the organ 

to that level. The following discussions of the Creede 

pinaceous fossils emphasize both preservational states of 

the fossils and comparative morphology of extant 

Pinaceae. 

Ovulate cones.-The 63 ovulate cone impressions 

from Creede display all degrees of abrasion and (or) 

deterioration prior to burial, a major factor not taken 

into account by Axelrod (1986, 1987). On plate 2 are 

arranged seven ovulate cones in order of increasing 

degree of abrasion. We arbitrarily designate these as 

abrasion stages 1 through 6; stage 1 shows little or no 

abrasion, but in stage 6 the cone is so seriously abraded 

that only the sturdy woody axis remains. In our opinion, 

cones of abrasion stage 4 + or greater are unidentifiable 

at the generic level. An assignment to a plus ( + ) 

designation means we consider the degree of abrasion to 

be intermediate between the illustrated examples. Using 

this method, for example, we support our assignment of 

the holotype of Pice a coloradensis Axelr. to Pinaceae 

Genus Indetermined by referring it to abrasion stage 5; 

the holotype of P. coloradensis is chosen to typify 

abrasion stage 5. Similarly, the holotype of Tsuga coloradensis Axelrod is referred to abrasion stage 4 + and is 

also transferred to the indetermined category (see pl. 2, 

figs. 5, 8). 

Examples of differential cone abrasion and the 

varying types and quality of fossil preservation are not 

unique to the Creede material. Illustrated here (pl. 3, 

figs. 1-3) are two early Pleistocene specimens from 

Potrero Canyon, California (Axelrod, 1967, 1990), and a 

Miocene impression from Turkey. Figure 1 is a naturally 

recovered longitudinal section of a cone that clearly 

exhibits all "abrasion" stages except the external surface 

impressions of stages 4 and 5. The specimen illustrated as 

figures 2 and 3 is a naturally occurring three-dimensional 

cone broken out of the rock matrix. Figure 2 shows the 

specimen with 10 attached cone scales that distinguish 

this as a pine. The outer surface of the cone scales 

represents approximately abrasion stage 2 + . When the 

small group of identifying cone scales is removed from 

the specimen (fig. 3), it becomes indetermined to genus; 

that is, abrasion stage 4 + (fig. 3). The basal part of the 

Turkish Miocene specimen (pl. 3, fig. 4), which has 

carbonized cone material present, would be impossible 

to assign to genus. In the apical area where the 

carbonized material is absent, the structure of the Pinus 

(Pinus) apophysis-umbo-mucro is clearly exposed. The 

differential preservational states illustrated by these 

fossils must be recognized in order to evaluate properly 

the taxonomic significance of the fossil record of 

Pinaceae. Taphonomic studies directed at understanding 

the processes and results of biological and (or) 

mechanical cone abrasion and deterioration are clearly 

needed in this aspect of paleobotany. An evaluation of 

the Creede ovulate cones based on abrasion criteria is 

given in table 1. 

Systematic Descriptions and Discussions 5 
Table 1. Abrasion stages of Creede ovulate cones of 

Pinaceae 

[Taxonomic assignment of pinaceous ovulate cones, including 

partial cones, as recognized by Axelrod (1987), the number censused 

(and illustrated) by Axelrod (1987), and their abrasion stages. One 

illustrated specimen of Pinus sanjuanensis (Axelrod, 1987, pl. 13, 

fig. 10) not in collections} 

Taxon 

Picea coloradensis 

P. lahontensis 

Pinus coloradensis 

P. crossii 

P. engelmannoides 

P. florissantii 

P. ponderosoides 

P. riogrande 

P. sanjuanensis 

Pseudotsuga glaucoides 

Tsuga petranensis 

Number 

5 (2) 

1 (2) 

2 (2) 

35 (2) 

1 (1) 

0 (1) 

2 (3) 

2 (3) 

1 (2) 

10 (4) 

1 (2) 

Abrasion stage 

5 and 6 

3 and4+ 

2 and4 

1 and2 

4 

2+ 

2 and4 

2 and4 

2 to 2+ 

4+, 5 and 6 

4+ and 5 

Winged Seeds.-Seeds that have wings are present 

in all the least modified members of the Pinaceae. Only 

certain members of Pinus, such as the nut pines 

(subsection Cembrae) and pinyon pines (subsection 

Cembroides ), have independently derived wingless seeds. 

Of the 10 genera in the family, only members of Pinus 

have developed a special condition in which the wing 

easily separates from the seed without any mechanical 

tearing of the wing. Such disarticulation of the wing from 

the seed is accomplished by the hygroscopic tissue of the 

wing. When the wing is dry a strip of wing tissue clasps 

firmly on either edge of the seed, holding the wing and 

seed together. In this condition, the wing and seed are 

said to be articulated but not adnate or fused. When such 

winged seeds become wet, the hygroscopic tissue adsorbs 

water and these cells swell, causing the two strips of wing 

tissue that clasp the seed to pull away from the seed. 

Articulated winged seeds have been independently 

derived in both subgenus Strobus and subgenus Pinus. 

The method of origin of the wing and seed in the 

Pinaceae is not widely recognized, but this is an 

extremely important feature because it relates to proper 

identification and the subsequent interpretation of the 

fossil record of winged seeds. The wing and seed in 

Pinaceae are of independent origin, unlike in Cupressaceae and Taxodiaceae, where the wing is derived from 

the outer coat of the seed. The wing in Pinaceae has its 

origin with the cone scale and will develop whether or not 

the ovule is fertilized and (or) matures. This feature has 

probably been independently recognized a number of 

times (James Eckenwalder, oral comm., 1985), but we 

6 Spermatopslda of Oligocene Creede Flora, Colorado 

are not aware of a more explicate discussion than the 

publication on Pseudotsuga menziesii (Mirb.) Franco by 

Allen and Owens (1972, p.120-123). If, as is typically the 

situation with the most basal and apical scales in 

Pinaceae, the scale is sterile or the ovule aborts early in 

development, the wing is developed without a normal 

seed and has a very different morphology than its winged 

seed sibling that has a fully developed wing and s_eed (pl. 

3, fig. 5). In fact, both fully developed and aborted types 

can be borne on the same cone scale (pl. 3, figs. 5e, g). 

Recognition of this attribute of Pinaceae wing/seed 

ontogeny is necessary because very differently appearing 

structures ensue, and these could be interpreted as 

different species or even different genera (pl. 2, figs. 10, 

11). For example, such undeveloped, atypical winged 

seeds in the Creede collections were distributed among 

four genera and seven species (Axelrod, 1987: Tsuga, pl. 

7, fig. 10; Picea, pl. 8, figs. 7-14; Pinus, pl. 9, figs. 5-7, pl. 

10, figs. 3, 6, and 7, and pl. 12, figs. 2-4; Pseudotsuga, pl. 

15, figs. 7, 8). When these undeveloped, atypical winged 

seeds are recognized, they should simply be assigned as 

atypical winged seeds of the genus, if possible, or 

otherwise as atypical winged seeds of Pinaceae. 

The winged seeds of Pinaceae form a significant 

part of the fossil record of the family. A literature 

review and personal observations of 15 paleofloras 

ranging in age from late Eocene to Pleistocene give the 

following relative abundance of the different types of 

detached organs found as Pinaceae impression fossils: 

defoliated axes, 0.28 percent; foliated axes, 0.85 percent; 

ovulate cones, 1.14 percent; detached ovulate cone 

scales, 1.73 percent; detached leaves and (or) fascicles, 

36.99 percent; winged seeds, 58.98 percent. Male cones 

are rare and not commonly recognized or recorded (see 

pl. 3, figs. 8, 9). The relatively high overall percentage of 

fossil winged seeds of Pinaceae (55-60 percent) indicates 

how important this record is to our understanding of the 

history of the family; however, based on the large number 

of misidentified winged seeds, the morphology of these 

organs is apparently the least understood feature in the 

family. For this reason, we present illustrated keys fot 

generic identification of the winged seeds of Pinaceae 

(the terminology we use for describing winged seeds is 

shown in figure 1; the keys are figures 2 and 3). Two keys 

are presented because single specimens of fossil winged 

seeds present only one surface of the seed and may not 

show the resin vesicles, pterostegium, or other features 

that are necessary for identification. The first key (fig. 2) 

is usable for living material and those fossils that have the 

adfacial surface of the winged seed exposed; such fossils 

should provide a high level of accurate identification, 

especially if well preserved. If the fossils expose the 
A. ORIENTATION (relative to position on cone scale) B. PARTS (of winged-seed) 

Wing (wing membrane or pteroderm) ----Admedial edge 

~~sy ~..,..,_---Abfacial surface 

Distal end 

Pterostegium 

Adfacial 

surface 

Abfacial 

surface 

Toward cone base 

Figure 1. Diagram showing proposed terminology for winged seeds of Pinaceae. 

abfacial surface or are of poorer quality preservation, 

more chance for error of identification exists. The 

greatest probability is to confuse small Pinus and Picea, 

and Tsuga-Larix-Pseudotsuga. The indentation on the 

admedial edge of Picea seeds distinguishes them from 

Pinus seeds. If the diagnostic characters of Tsuga (resin 

vesicles and wing cellular pattern), Larix (convex adfacial 

surface of seed and wing cellular pattern) and Pseudotsuga (flattened adfacial surface of seed and wing cellular 

pattern) are poorly preserved or do not show because of 

orientation in the rock, it would be more realistic and 

conservative to refer such specimens to Pinaceae cf. 

Tsuga-Larix-Pseudotsuga. These types of fossils can also 

be compared to other well-preserved specimens from the 

same fossil flora, giving tentative assignments based on 

sharing of less critical characters. Whatever course is 

taken, we suggest that poorly preserved material should 

not be "overworked" by basing new species on such 

material. 

The winged seeds of Pinaceae from Creede 

(excluding Abies) are keyed out as follows. 

A. Wing adnate 

1. Admedial edge of wing 1 em long or less 

a. Seed small, < 3 mm ................................. atypical Pinus 1 

b. Seed large, > 3 mm 

1. Admedial edge of proximal end of seed with 

concave indentation ...................................... .Picea 

2. Concave indentation absent ........ atypical Pinus 2 

2. Admedial edge of wing > 1 em long 

a. Seed small, < 3 mm long ........................ atypical Pinus 3 

b. Seed intermediate, 3-5 mm long ............... Pinus type A 

c. Seed large, > 5 mm long ............................. .Pinus type B 

B. Wing disarticulated 

1. Admedial edge of wing < 2 em long 

a. Vacated seed area < 1.5 mm wide ....... atypical Pinus 4 

b. Vacated seed area 2.5-3.5 mm wide ....... .Pinus type C 

2. Admedial edge of wing > 2.5 em long, vacated seed area 

> 5 mm wide .................................................. Pinus type D 

I. Seeds with resin vesicles; part of abmedial cellular pattern of wing directed toward abmedial 

edge of wing, intersecting with edge at an 

angle 

A. Pterostegium well developed on adfacial 

surface of seed 

1. Wing widest in distal half .... Abies 

2. Wing widest in proximal half 

a. Wing 1 1 I 4-1 1 /2 length of seed 

. . . . . . . . . . . . . . . . . . . . . . . Keteleeria 

b. Wing 2 1/2-3 1/4 length of seed 

. . . . . . . . . . . . . . . . . . . . . Pseudolarix 

B. Pterostegium little (or not) developed on 

adfacial surface of seed 

1 . Wing widest in distal half . . . Cedrus 

2. Wing widest in proximal half ... Tsuga 

II. Seeds without resin vesicles 

A. Cellular pattern of wing undulatory and 

parallel with long axis of wing; wing typically 

constricted or wrinkled orthogonal to long axis 

of wing ........................... Pinus 

B. Cellular pattern of wing not as in A 

1 . Wing widest in distal half; seed typically 

with indentation at proximal end of admedial edge; pterostegium typically not 

expressed . . . . . . . . . . . . . . . . . . . . . Picea 

2. Wing widest in proximal half; small 

area of pterostegium on proximal end of 

seed 

a. Seed distally rounded, adfacial 

surface convex . . . . . . . . . . . . . Larix 

b. Seed distally angular, adfacial 

surface flattened ..... Pseudotsuga 

Figure 2. Illustrated key for generic identification of winged 

seeds of Pinaceae that have adfacial surface exposed. Cathaya 

was not available for study. 

Systematic Descriptions and Discussions 7 
The total winged seeds assigned to each species as 

recognized by Axelrod (1987, tables 1-4) and their keyed 

equivalents follow. 

Abies concoloroides 

A. rigida 

Picea coloradensis 

P. lahontensis 

P. sonomensis 

Pinus alvordensis 

P. coloradensis 

P. crossii 

P. macginitiei 

P. ponderosoides 

P. riogrande 

P. wasonii 

Pseudotsuga glaucoidescoides 

Tsuga petranensis 

2 (not in collections, not evaluated) 

30=Abies rigida ( + Picea 

lahontensis in part) 

10=atypical Pinus 1 

16 =Picea sp. ( + Abies rigida, 

atypical Pinus 2, indet. Pinaceae 

in part) 

2 = atypical Pinus 1 and 3 

6 =atypical Pinus 1 and 3 

3 =atypical Pinus 2, 4, and Pinus 

type A 

147 =Pinus type C 

1 =Pinus type D 

22 =Pinus type C 

None censused but 3 figured = 

Pinus type A and atypical Pinus 4 

At least 3 = Pinus type B 

4 = atypical Pinus 2 

2 = atypical Pinus 2 

The reevaluation just listed, concomitant with our 

systematic revisions, indicates that pinaceous winged 

seeds from Creede can be enumerated as follows: 

Abies rigida 31 

Picea sp. 9 

Pinus crossii 172 

P. sp. 2 1 

P. spp. indet. 33 

Fascicles and isolated needles.-Two types of 

fascicles occur at Creede: a two-needled morphotype that 

has leaves 2-3 em long (type 1), and a five-needled 

morphotype that has leaves varying from slightly more 

than 2 to about 6 em long (type 2). Both types have a 

deciduous sheath and shoot morphology typical of the 

soft (Haploxylon) pines of subgenus Strobus. By shoot 

morphology we are referring to the shape and structure 

of the basal "short shoot" axis to which the leaves are 

attached. In the soft pines, the shoot is circular in outline, 

and because the sheath is deciduous, the shoot has small 

scars exposed where the individual sheath elements were 

attached to the shoot. In the Creede material, the 

exposed shoot appears as a shortened cylinder that is 

approximately as wide as tall. The hard pines (subgenus 

Pinus) have a more pronounced, bilaterally symmetrical, 

slightly bulbous shoot, and the sheath is persistent 

(except in Pinus leiophylla Scheide & Deppe; W. B. 

Critchfield, oral commun., 1980). Mechanical removal of 

the persistent sheath very typically results in individual 

leaves falling off. 

8 Spermatopsida of Oligocene Creede Flora, Colorado 

I. Cellular pattern of wing undulatory and parallel 

with long axis of wing; wing constricted or 

wrinkled orthogonal to long axis of wing 

................................... Pinus 

II. Cellular pattern of wing not as in I 

A. Part of abmedial cellular pattern of wing 

directed toward abmedial edge of wing, intersecting edge at angle 

1. Wing widest in distal half 

a. Proximal end of seed blunt; seed 

oblong, oval, or triangular ... Abies 

b. Proximal end of seed not blunt; 

seed typically oblong ... '. . . Cedrus 

2. Wing widest in proximal half 

a. Wing + seed typically less than 

2 em long ................ Tsuga 

b. Wing + seed typically greater than 

2 em long; wing tissue very dense 

along abmedial edge of seed 

1) Wing 1 1/4-1 1/2 length of 

seed. .. . . . . . . . . . . . . Keteleeria 

2) Wing 2 1 /2-3 1 I 4 length of 

seed . . . . . . . . . . . . . Pseudolarix 

B. All cellular pattern of wing parallel to long 

axis of wing; no constriction or wrinkling 

1. Wing widest in distal half; seed typically 

with indentation at proximal end of admedial edge . . . . . . . . . . . . . . . . . . . Picea 

2. Wing widest in proximal half 

a. Seed distally rounded . . . . . Larix 

b. Seed distally angular .. Pseudotsuga 

~ 

~ 

~\\ 

\)) 

~~ Figure 3. Illustrated key for generic identification of winged 

seeds of Pinaceae that have abfacial surface exposed. 

Cathaya was not available for study. 

Variation in needle length of the five-needled type 

2 is atypically great for extant pines. A histogram of 

needle length (fig. 4) indicates that two taxa could be 

represented, one that has needle length less than 3.5 em 

(dominantly 2.0-3.0 em) and one that has needle length 

more than 4.0 em. One shoot (pl. 13, fig. 4) however, has 

17 attached fascicles in which needle length varies from 

2.4 to 6.0 em; the long-needled fascicles are both at the 

basal and at the apical parts of the incomplete shoot, and 

the short-needled fascicles are in between. These 

relations could be interpreted as the two groups of 

long-needled fascicles representing the moister, earlier 

parts of two growing seasons and the short -needled 
fascicles representing the drier, later part of the growing 

season in between. This interpretation is consistent with 

the dry summers inferred for the Creede assemblage 

(Wolfe and Schorn, 1989). 

The two types of Creede fascicles were distributed 

among six species, five subsections, and two subgenera 

(Axelrod, 1987). The absence of a sheath on any of the 

Creede specimens immediately eliminates their 

suggested assignment to two of the subsections of subgenus Pinus. As determined and censused by Axelrod 

(1987) and the morphotypes represented, the Creede 

pine fascicles are: 

Pinus alvordensis 

P. coloradensis 

P. crossii 

P. jlorissantii 

P. sanjuanensis 

P. wasonii 

17 (type 1) 

165 (type 2) 

703 (type 2) 

535 (type 2) 

38 (type 1) 

16? (type 1) 

Based on our systematic revisions and Axelrod's (1987) 

census data, fascicles of Pinus crossii (type 2) are 

abundant, but fascicles of P. sanjuanensis (type 1) are 

uncommon. 

Three isolated needles were assigned to Pseudotsuga (Axelrod, 1987, p. 96). Extant Pseudotsuga has 

needles that have a base constricted to form a distinct 

petiole, have marginal resin canals, and are planar; none 

of these features are discernible on the Creede 

specimens. These Creede leaves have a triangularshaped leaf base and cross section and are approximately 

2.5 em long; they probably represent individual leaves 

that were separated from fascicles of Pinus crossii, 

although we conservatively relegate such specimens to 

the indetermined category. 

Foliated axes.-Foliated axes, other than those of 

Juniperus and Pinus, were assigned to two species of 

Abies and one species of Pseudotsuga (Axelrod, 1987). 

(/) 

z 

w 

~ 20 u 

w 

a.. 

(/) 

LL 

0 

Iz 

w 10 u 

a: 

w 

a.. 

o~~~~~--~~~~~~~~==~~ 

2.0 2.5 3.0 3.5 4.0 4.5 5.0 

FASCICLE LENGTH, IN CENTIMETERS 

Figure 4. Histogram of needle length for five-needled 

fascicles of Creede Pinus. Based on 79 specimens. 

Because none of these axes have leaves that have 

constricted leaf bases that form a petiole, none can 

represent Pseudotsuga; the specimens assigned to 

Pseudotsuga have circular leaf bases and (or) leaf scars 

typical of Abies (pl. 1, fig. 8). 

Foliated axes that have longer, more flattened, 

pseudodistichously arranged leaves were referred to 

Abies concoloroides (except for those referred to 

Pseudotsuga). Foliated axes that have falcate-shaped 

leaves that appear crowded on the upper sides of the axes 

were referred to A. rigida. Such morphologic differences, 

however, can occur in branches of the same tree. For 

example, Sudworth (1908, p. 125) noted that in A. 

amabilis (Loud.) Forbes, "The leaves of lower crown 

branches are flat** *They are about 1 Y4 inches long, and, 

by a twist in the bases of those on the lower sides of the 

branches, they appear massed on the top sides [pseudodistichous]. Branches of the uppermost part of the crown 

have shorter and stouter leaves, about three-fourths of an 

inch long*** and stand erect in dense masses on the tops 

of the sprays." Sudworth (1908) and Liu (1971) also 

described similar conditions for A. concolor (Gord.) 

Parry, A. procera Rehd., and A. magnifica Murr. The 

dimorphic condition in the Creede axes of Abies is thus 

similar to that in single species of extant Abies, and we 

consider all these foliated axes to represent A. rigida. 

Genus ABIES Miller 

Abies rigida Knowlton 

PL-.te 1, figures 3--10 

Abies rigida Knowlton, 1923, p. 186, pl. 41, figs. 1, 2, 4, 5. Axelrod, 1987, p. 83, pl. 

5, figs. 1-8; pl. 6, figs. 1-10. 

Abies longirostris Knowlton, 1923, p. 187, pl. 42, figs. 1, 2. 

Abies concoloroides auct. non R. W. Brown. Axelrod, 1987 (part), p. 82, pl. 7, figs. 

1-3, 6. 

Picea lahontensis auct. non MacGinitie. Axelrod, 1987 (part), p. 85, pl. 8, fig. 2. 

Pseudotsuga glaucoides Axelrod, 1987 (part, nontypic), p. 96, pl. 15, figs. 1, 3, 

9, 10. 

Wolfe and Schorn (1989) designation: Abies rigida. 

Description. -Foliated axis more than 5.6 em long, 

0.5-0.6 em wide, circular leaf scars slightly less than 0.1 

em wide, almost flush with surface of axis; about 20 

leaves preserved, about 1.5 em long and slightly more 

narrow than leaf scars; leaf shape falcate, base. a circular 

expansion slightly wider than leaf, apex rounded, leaves 

twisted so as to appear to be derived from and directed 

away from one side of axis (presumably adaxial side); 

other axes have distichously arranged leaves that are 

about 6 em long. Ovulate cone scale at least 1-2 em wide, 

1.0-1.5 em long (scale and stalk); scale-length to stalklength ratio about 2:1; scale shape cuneate to reniform; 

bract scale exserted, length relative to cone scale 

variable, as much as 5.0 em long; width variable, 

commonly widest under cone scale (where laminar tissue 

Systematic Descriptions and Discussions 9 
is expanded but no lobes formed, tapering to a stiff, rigid 

apex. Winged seeds have wing-length to seed-length ratio 

of about 1:1; seeds typically ovate to obovate, seed-length 

to pterostegium-length ratio about 1.25:1. 

Discussion.-The detatched cone scales and 

foliated axes of the Creede fir were originally described 

as two distinct species (Knowlton, 1923); however, Axelrod (1987) validly recognized their equivalency and 

combined the two different organs into a single species. 

Included in Abies rigida are ovulate cone scales (attached 

bract scales are present or absent), winged seeds, axes 

(foliated or defoliated), and isolated leaves. 

The long exserted bract scale is absent from two 

Creede cone scales assigned to Abies concoloroides R. W. 

Br. (Axelrod, 1987, pl. 7, figs. 2, 3). The size and shape of 

these cone scales, the ratio of stalk length to scale length, 

and the shape and relative size of the wing impressions on 

the scales are all characters shared, however, with cone 

scales assigned to A. rigida. 

One winged seed assigned to Picea lahontensis 

(Axelrod, 1987, pl. 8, fig. 2), unlike seeds of extant Picea, 

has no indentation on the proximal end of the admedial 

edge of the seed. The shape of the seed and the wing, the 

faint cellular pattern of the wing, and the arrangement of 

the seed-wing juncture are characters found in winged 

seeds of Abies rigida. 

Numerous unfoliated and foliated axes of Abies are 

recognizable by, respectively, the presence of circular leaf 

scars and the absence of leaf bases constricted into a 

petiole. Knowlton (1923, p. 187) suggested the possibility 

that two species of fir were represented by foliated axes 

in the Creede flora. Axelrod (1987) also recognized two 

morpho types of Abies axes, assigning the most common 

type to A. rigid a and a type that has pseudodistichous leaf 

arrangement to A. concoloroides; however, some axes 

that have a pseudodistichous leaf arrangement and 

unconstricted leaf bases (and thus represent the second 

Abies morpho type) were invalidly assigned by Axelrod 

(1987) to Pseudotsuga. As previously discussed, a 

condition in which leaves are flattened into a pseudodistichous arrangement is common on lower branches of 

extant firs, even those Abies that have leaves of the upper 

branches twisted and directed toward the upper (adaxial) 

side of the branch (Sudworth, 1908; Liu, 1971). The 

variation displayed by extant firs supports assignment of 

all these foliated axes from Creede to a single entity. 

Relative to leaf length, the specimens assigned by Axelrod (1987) toA. concoloroides,A. rigida, andPseudotsuga 

also overlap (fig. 5). 

The cone scales of Abies rigida were, because of the 

highly exserted sub tending bracts, originally compared to 

the extant A. bracteata D. Don (Knowlton, 1923). This 

suggested relationship may be valid. The extant genus 

most closely related to Abies is Cedrus, in which the bract 

is reduced and adnate to the cone scale. This relation 

1 0 Spermatopsida of Oligocene Creede Flora, Colorado 

U!~Doo\ RL00 

2 3 

121 [7J 

LEAF LENGTH, IN CENTIMETERS 

~ Abies·"concoloroides" 

D Abies rigida 

[ZJ "Pseudotsuga glaucoides" 

Figure 5. Histogram of leaf length in Creede Abies. 

could suggest that reduced, inserted subtending bracts, 

as found in many extant species of Abies, represent the 

plesiomorphic condition. If this suggestion is valid, then 

the highly exserted bracts of A. rigida and A. bracteata 

could represent a synapomorphy and indicate that the 

two taxa are more closely related to one another than to 

other species of Abies. Until a thorough cladistic analysis 

of extant Abies is undertaken, however, such suggested 

relationships remain speculative. 

That the foliage of Abies rigida has little similarity 

to that of A. bracteata (Axelrod, 1987) may not be 

significant relative to phyletic relationships. The extant A. 

bracteata inhabits far milder climates than the climate 

inferred for A. rigida (Wolfe and Schorn, 1989); foliage of 

many extant Abies that inhabit subalpine environments 

have, as in A. rigida, leaves twisted and directed toward 

the upper side of the branch. 

Two winged seeds from Creede were attributed to 

Abies concoloroides R. W. Br. (Axelrod, 1987, pl. 7, figs. 

4, 5). Determination of the taxonomic status of these 

seeds, which were not placed in the University of 

California Museum of Paleontology, must await their 

relocation and examination. A. concoloroides is a poorly 

understood concept, however, and fossil material that 

appears to be related to the extant A. concolor (Gord. & 

Glend.) Lindl. should not indiscriminately be assigned to 

A. concoloroides. Abies concoloroides was founded 

(Brown, 1940) on an impression of the abaxial surface of 

an ovulate cone scale that does not have a bract scale and. 

on a foliated axis of a Picea (see later discussion of P. 

lahontensis); however, because A. concoloroides has 

never been formally described or diagnosed, the epithet 

must be considered an illegitimate name. The foliated 

axes and ovulate cone scales from Creede attributed to A. 

concoloroides (Axelrod, 1987) are reassigned toA. rigida. 

Lectotype.-USNM 36505. 

Syntypes.-USNM 36507-36509. 

Hypotypes.-VSNM 36517, 36518; UCMP 

7248-7263, 7265-7281, 7283-7286, 7308, 7381, 7384, 

7410-7416, 7426-7430. 
Genus PICEA Dietrich 

Picea sp. 

Plate 3, figure 6 

Picea lahontensis auct. non MacGinitie. Axelrod, 1987 (part), p. 85, pl. 8, figs. 

4-6. 

Wolfe and Schorn (1989) designation: Picea sp. 

Discussion. -Of the three species of spruce 

recognized by Axelrod (1987) in the Creede flora, only 

one is considered validly to represent Picea. The Creede 

spruce is part of a big-seeded complex that includes P. 

lahontensis Macg. and P. magna MacG .; this complex is 

no longer extant in North America and may represent 

one or more lineages. From the latest Eocene P. magna 

to the Neogene P. lahontensis, morphologic trends are 

for the winged seed to increase in size and for the wing to 

become longer relative to the seed. The Creede spruce 

seeds are most similar to the types of P. magna but differ 

in having shorter and broader wings; these differences 

are not morphologically intermediate between P. magna 

and P. lahontensis and suggest that the Creede Picea, in 

retaining an apparently more primitive morphology, 

could be a sister lineage to the P. magna-P.lahontensis 

lineage. A complete morphometric analysis of these 

big-seeded taxa of spruce should further resolve their 

relationships. 

One winged seed assigned to Picea lahontensis 

(Axelrod, 1987, pl. 8, fig. 2) represents Abies rigida. 

Characteristic of spruce seeds relative to other members 

of Pinaceae is a concave indentation on the proximal end 

of the admedial edge of the seed, and this indentation is 

absent in the seed just cited. The fossil also has a cellular 

pattern on the wing directed outward toward the abmedial edge of the wing; this feature is restricted to, and 

diagnostic of, the abietoid Pinaceae (Abies, Cedrus, Keteleeria, Pseudolarix, and Tsuga). Some other supposed 

spruce seeds (Axelrod, 1987, pl. 8, figs. 7-14) are 

considered to represent atypical seeds of pine (see later). 

None of the Creede ovulate cones assigned to Picea 

(Axelrod, 1987) have characters diagnostic of the genus. 

Except for one specimen, all these cones are too abraded 

for generic identification. The taxonomic disposition of 

these cones follows. 

Picea lahontensis (Axelrod, 1987, pl. 8, fig. 1) = 

Pinaceae genus and species indetermined. This cone 

(and others of abrasion stage 4 + or greater) exhibits 

no features that are diagnostic to genus. The presence 

of spirally arranged, woody, persistent cone scales (or 

their remnants) indicate that the cone belongs to a genus of Pinaceae that has persistent cone scales (Cathaya, Larix, Keteleeria, Picea, Pinus, Pseudotsuga, 

Tsuga). The size and shape of the cone suggest Keteleeria, Picea, Pinus, and perhaps Pseudotsuga. Because other, better preserved pinaceous cones in the 

Creede collections primarily represent Pinus crossii, 

the cone most probably represents that taxon. Other 

supposed spruce cones (Axelrod, 1987, pl. 8, fig. 15; 

pl. 8, fig. 16 [holotype of Picea co/oradensis Axelr.]) 

represent abrasion stages 6 and 5, respectively; these 

may also represent Pinus crossii. 

Picea lahontensis (Axelrod, 1987, pl. 8, fig. 3) = Pinus 

ct. P. crossii Knowlt. This cone fragment (a fragment 

cited as homeotype UCMP 7315 is part of the same 

cone; see our pl. 2, fig. 3) has the dorsal umbos characteristic of Pinus. The cone represents abrasion 

stage 3. 

The single defoliated axis assigned to a spruce 

probably represents an angiosperm. 

Specimens.- UCMP 7302, 7304-7307,7310,7313. 

Genus PINUS Linnaeus 

Subgenus STROBUS Lemmon 

Section PARRYA Mayr 

?Subsection CEMBROIDES Engelmann 

Pinus sanjuanensis Axelrod 

Plate 1, figures 11, 12 

Pinus sanjuanensis Axelrod, 1986, p. 613, figs. 12-16. Axelrod, 1987 (part), p. 95, 

pl. 13, figs. 2-8, 10, 11. 

Pinus alvordensis auct. non Axelrod, 1944. Axelrod, 1986 (part), p. 627, figs. 

79-81. 

Wolfe and Schorn (1989) designation: Pinus sp. 

Description. -Ovulate cone . uncertain (specimen 

misplaced, and detailed description awaits relocation of 

specimen). Fascicles have two leaves, 1.2-3.1 em long, 

mean length of 36 fascicles (including leaf and short 

shoot) 1.6 em, about 0.1 em wide, sheath deciduous. 

Discussion-Pinus sanjuanensis was considered 

(Axelrod, 1986, 1987) to be most similar to the extant 

pinyon, P. edulis Engel. The ovulate cone that is the 

holotype of the species has not been located. The 

illustration of the cone lacks details sufficient to make a 

satisfactory evaluation of the specimen, and the formal 

description is similarly unsatisfactory. Close affinities of 

this cone are thus speculative, but we will follow Axelrod 

(1986, 1987) and consider this cone that of a pinyon. 

Had the terminology formulated by Klaus (1980) 

(fig. 6) been consistently employed, the affinities of Pinus 

sanjuanensis could be deciphered. This species is 

described, however, as "***apophyses subdued; umbo 

mucronate, centro in position on scales***" (Axelrod, 

1986, p. 613). Klaus (1980, fig. 4) used traditional 

terminology in reference to either terminal or dorsal 

position of the umbo relative to the apophysis. In cones 

that have dorsal umbos, Klaus further differentiated the 

position of the mucro (spine, thorn, or prickle): (1) 

mucro positioned on the transverse keel ( centromucronate) and (2) mucro positioned distally above the 

transverse keel in the upper field of the umbo ( excentromucronate ). Does Axelrod's "centro in position on the 

scales" refer to the umbo as being centrally positioned on 

the apophysis (that is, dorsal) or to the mu~ro being 

centrally positioned on the umbo ( centromucronate )? 

Although the affinities of the holotype of Pinus 

sanjuanensis are somewhat uncertain, we have, as did 

Axelrod (1987), included two-needled fascicles in this 

Systematic Descriptions and Discussions 11 
a 

b 

c 

d 

Figure 6. Drawing showing general terminology of apophysis, umbo, and mucro of ovulate cone scale in Pinus. a, 

upper umbo field; b, transverse keel; c, mucro (centroerect); d, lower umbo field. Modified from Klaus (1980). 

species so as not to proliferate nominal species. We have 

also included morphologically indistinguishable fascicles 

assigned by Axelrod (1987) to P. alvordensis. Axelrod 

(1986, p. 627) noted similarities between the fascicles he 

assigned to P. sanjuanensis or to P. alvordensis but 

differentiated fascicles of P. sanjuanensis on the basis 

that "***the fascicle sheath is smaller as compared with 

those of P. alvordensis ." None of these two-needled 

fascicles have sheaths, which were presumably 

deciduous. 

Subsection Contorlae, to which Pinus a/vordensis 

was assigned, has persistent sheaths, and subsection 

Cembroides, to which P. sanjuanensis was assigned, has 

deciduous sheaths. Perhaps Axelrod was referring to the 

short shoot, upon which the leaves are borne, as the 

"fascicle sheath," but, if so, the fascicles referred to P. 

sanjuanensis are still inseparable from those referred to 

P. alvordensis. 

Pinus sanjuanensis was considered (Axelrod, 1986) 

to be related toP. ballii R. W. Br. from the Eocene Green 

River flora. Brown (1934) described P. ba/lii on the basis 

of a single fascicle that he, as well as subsequent authors 

(MacGinitie, 1969; Axelrod, 1986, 1987), assumed to 

have three leaves. Four leaves are visible on the 

specimen, the fourth tip being observable just above the 

tip of the leaf to the right. This specimen, however, is 

poorly preserved and is an inadequate basis for the 

supposed earliest record of pinyon pines. MacGinitie 

(1969) added a supposed cone to P. ballii, an addition 

accepted by Axelrod (1986, 1987). This open "cone" has 

"scales" that appear to arise from a common point of 

origin; the scales are narrow, have no indication of large 

seed cavities that would be present in a pinyon pine, and 

have no obvious apophysis-umbo structures at the distal 

ends. This "cone" certainly does not represent Pinus and 

most probably is an angiosperm fructification. 

Holotype.-VCMG 19701. 

Paratypes.-UCMP 7239-7244, 7247, 7393, 7400. 

Hypotypes.-VCMP 7173-7176,7236-7238,7396, 

7398, 7399. Note that specimen numbers were reversed 

on UCMP 7235 and 7174; UCMP 7174 was first 

12 Spermatopsida of Oligocene Creede Flora, Colorado 

illustrated as Pinus alvordensis (Axelrod, 1986, fig. 79), 

and then a second specimen was assigned the same 

number and also illustrated asP. alvordensis (Axelrod, 

1987, pl. 3, fig. 3). This second specimen, which is a 

fascicle of P. crossii that has three leaves missing, is now 

assigned UCMP 7235. 

Subsection BALFOURIANAE Shaw 

Pinus crossfi Knowlton 

PL.'lte 2, figures 1, 2; pL.'lte 3, figures 7-13; plate 13, figure 4; text figures 7A, B 

Pinus crossii Knowlton, 1923, p. 185, pl. 41, figs. 3, 8-10. Bailey, 1970, p. 239, figs. 

34-37. Axelrod, 1986, p. 73, figs. 34-42. Axelrod, 1987, p. 91, pl. 7, figs. 

11-16; pl. 9, figs. 10-13; pl. 13, figs. 9, 12. 

Pinus co/oradensis Knowlton, 1923, p. 186, pl. 41, fig. 6. Axelrod, 1986, p. 615, 

figs. 43, 48 51, 53. Axelrod, 1987, p. 90, pl. 10, figs. 1, 2. Axelrod, 1986 (part), 

p. 615, figs. 48, 51, 53. 

Pinus simi/is Knowlton, 1923, p. 186, pl. 41, figs. 11, 12. 

Pinus aristata fossilis Cockerell, 1934, p. 573, fig. 1. 

Pinus ponderosoides Axelrod, 1986 (part, typic), p. 623, figs. 64-66. Axelrod, 1987 

(part), p. 94, pl. 11, figs. ?-7, 9, 10. 

Pinus wasonii Axelrod, 1987, p. 96, pl. 14, figs. 6-12. 

Pinus a/vordensis auct. non Axelrod, 1944. Axelrod, 1987 (part), p. 89, pl. 9, fig. 

3. 

Pinus florissantii auct. non Lesquereux. Axelrod, 1986 (part), p. 607, fig. 9. 

Axelrod, 1987, p. 92, pl. 14, figs. 1-5. 

?Pseudotsuga glaucoides Axelrod, 1987 (part, nontypic), p. 96. 

Wolfe and Schorn (1989) designation: Pinus crossii. 

Description. -Ovulate cone shape oblong to ovate, 

typically elliptic; as much as 10 em long, 4.5 em wide; 

apophysis in plane or facial view about 1.4 em wide, 1.0 

em high, width to height ratio about 1.5:1; umbo dorsal; 

mucro centroerect, as much as 0.8 em long, shape 

rhombic, protruding above plane of apophysis. Winged 

seed disarticulated (rarely found articulated); vacated 

seed area about 0.3 em wide (rare seeds 0.7 em long); 

line drawn along admedial edge of wing passes through 

admedial half of vacated seed area; seed wing has admedial edge about 1.2-1.5 em long, about 0.6 em wide; 

abmedial edge tapering distally, distal end typically 

rounded to obtuse. Fascicles have five (some four) 

leaves; typically about 2.5-4.5 em long but ranging from 

1.6 to 6.0 em, 0.1 em wide; slightly curved; sheath 

deciduous. 

Discussion.-As circumscribed here, Pinus crossii 

Know It. is the common pine of the Creede flora. Using 

Axelrod's (1987) census data and our reassessment of 

the species, P. crossii is represented by approximately 40 

ovulate cones, 170 winged seeds, and 1,400 fascicles. 

Most severely abraded ovulate cone remnants and 

atypical winged seeds probably also belong to this taxon. 

Pinus crossiiwas originally founded on five-needled 

fascicles that Knowlton (1923) clearly recognized as a 

soft pine because of the deciduous sheath. Axelrod 

(1986) gave a lucid account of our historical 

understanding of the species and the morphologic 

evidence that relates this fossil species to the extant P. 

aristata Engelm. of the southern Rocky Mountains of 
Colorado and New Mexico. Although Axelrod 

considered the fossil species to be allied to P. aristata, he 

(1986, p. 615) also recognized that "***there are 

sufficient differences to indicate that P. crossii probably is 

ancestral to the living bristlecone pine." 

Identification of impression fossils of ovulate Pinus 

cones involves some interpretation regarding the threedimensional reconstruction of the apophysis-umbomucro relations. The horizontal pressure during diagenesis of the sediments is minimal and the horizontal 

outline of the cone is basically the original outline. The 

vertical pressure, however, significantly distorts original 

relations, and casts made from natural molds of 

impressed specimens must be interpreted. The place to 

observe least distortion of the original relations is along 

the margins of the cone impression where the cone scale 

can be seen in profile; care must be taken not to confuse 

the obliquely oriented apophyses with profiles of the 

cone scales themselves. Profile and plane view 

reconstructions of the three types (excluding P. sanjuanensis, the sole cone of which is unavailable) of ovulate 

pine cones from Creede are illustrated in figure 7. 

Knowlton originally described Pinus coloradensis 

from a single, incomplete ovulate cone impression. He 

noted (1923, p. 186) that the "***scales [are] much 

thickened at the end***" and that the impression was 

missing "***a few scales at the base***" Axelrod (1986, 

1987), in adding two cone impressions to this species, 

contended (1986, p. 617) that the cone scales were thin 

rather than thickened. Axelrod's illustrations (1986, figs. 

43, 44 ), including that of a latex cast, of Knowlton's type 

show, however, that the profile of the scales has a 

thickened apophysis and protruding umbo (see our fig. 

1B, which is a specimen referred by Axelrod, 1987, toP. 

coloradensis ). Only in plane view do the cone scales 

Figure 7. Drawing showing ovulate cones of Creede 

species of Pinus. Plane and profile views. A, 8, P. crossii 

Knowlt. C, P. riogrande Axelr. D, P. sp. 1. 

appear thin, but this resulted from vertical compression. 

Thus, the type and one of Axelrod's (1987) unfigured 

ovulate cones of P. coloradensis are referred to P. crossii. 

The third specimen referred toP. coloradensis (Axelrod, 

1986, fig. 45; 1987, pl. 10, fig. 4) is the apical part of a 

cone that is abraded to stage 4 and is transferred to P. cf. 

P. crossii. 

The holotype of Pinus ponderosoides Axelr. is also 

placed in P. crossii. This specimen is the lateral portion of 

an ovulate cone that has 16 complete or partially 

complete apophyses. The apophyses are as high as 

1.0-1.2 em and are slightly wider, giving a length to 

height ratio of almost 1.5:1. The dorsal umbo protrudes 

distinctly beyond the plane of the apophysis and is 

centromucronate; the mucro is erect. The full mucros are 

missing and are inferred to be abraded; this is the typical 

situation in P. crossii cones. 

The disarticulated seed wings and an ovulate cone 

scale originally included in Pinus ponderosoides are also 

considered to represent P. crossii. Two strongly abraded 

ovulate cones referred to P. ponderosoides (Axelrod, 

1987) are placed in P. cf. P. crossii. 

The syntypes of Pinus simi/is were synonymized 

under P. florissantii, to which was added a single Creede 

ovulate cone (Axelrod, 1986, 1987). This cone (pl. 2, fig. 

2; abrasion stage 2) has dorsal umbos and represents P. 

crossii of subsection Balfourianae, in contrast to the type 

of P. florissantii, which has terminal umbos and thus 

represents a soft pine of the P. flexilis alliance in 

subsection Strobi (Axelrod, 1986). 

A few moderately large winged seeds could be 

assigned to subsection Strobi based on their adnate seeds; 

however, considering the abundance of the typically 

disarticulated wings of Pinus crossii, a few specimens in 

which seed and wing failed to detach might be expected. 

The Creede five-needled fascicles that have 

deciduous sheaths were segregated by Axelrod (1986, 

1987) into four taxa: Pinus florissantii, P. coloradensis, P 

crossii, and the newly institutedP. wasonii. We can see no 

morphologic basis for such specific distinctions and 

consider that all these fascicles most probably represent 

the same biologic entity. Knowlton (1923, p. 186) 

considered that the leaves of P. simi/is were 

indistinguishable from P. hambachii Kirchn. from Florissant. An enlargement of the negative of the type of P. 

hambachii (illustrated at natural size by MacGinitie, 

1953, pl. 19, fig. 1) indicates that the specimen is the 

frayed end of a reedlike plant. 

Some unfigured homeotypes of Pseudotsuga glaucoides (Axelrod, 1986, p. 96) probably represent 

detached leaves of Pinus crossii. 

Lectotype.-USNM 36514. 

Syntypes.-USNM 36511-36513. 

Systematic Descriptions and Discussions 13 
Hypotypes.-VCMG 34154; UCMP 7181-7185, 

7190-7198, 7200-7202, 7205-7208, 7212-7214, 7216, 

7218-7222, 7235 (see UCMP 7174 under Pinus sanjuanensis) 7246, 7317, 7326-7328, 7330-7335, 7338, 7437, 

7339-7342, 7344-7352, 7354, 7358 (counterpart of 

7354), 7362, 7368, 7380, 7387, 7390-7392, 7401-7406, 

7408, 7409, ?7431-7433, 7862-7866; USNM 36506, 

36516. 

Pinus cf. P. crossii Knowlton 

Plate 2, figures 3, 4 

Pinus engelmannoides Axelrod, 1986, p. 621, fig. 55. Axelrod, 1987, p. 92, pl. 9, 

fig. 8. 

Picea lahontensis auct. non MacGinitie. Axelrod, 1987 (part), p. 85, pl. 8, fig. 3. 

Pinus coloradensis auct. non Knowlton. Axelrod, 1986 (part), p. 615, fig. 45. 

Axelrod, 1987 (part), p. 90, pl. 10, fig. 4. 

Pinus florissanti? auct. non Lesquereux. Knowlton, 1923, p. 186, pl. 41, fig. 7. 

Pinus ponderosoides auct. non Axelrod, 1986. Axelrod, 1987 (part), p. 94, pl. 11, 

figs. 1, 2. 

Pinus riogrande Axelrod, 1986 (part, nontypic), p. 624. Axelrod, 1987 (part), p. 

94. 

Wolfe and Scorn (1989) designation: Pinus crossii. 

Discussion.-All these specimens are ovulate 

cones of Pinus. Most are abraded to approximately stage 

4; they cannot be identified to subgeneric level, and none 

are worthy of standing as holotypes. The material shows 

the size and general apophysis-umbo shape of P. crossii 

and most probably represents the abundant Creede pine. 

Some might, if better preserved, prove to represent a 

different entity than P. crossii, but this is speculative. Of 

the approximately 50 Creede cones of Pinus that are 

determinable to species, almost all represent P. crossii, 

and probably almost all the specimens just cited also 

represent this taxon. 

Specimens.- UCMG 27381; UCMP 7211, 7315, 

7316 ( = apical third of UCMP 7315), 7230, 7353, 7357; 

USNM 36510. 

Subgenus PINUS 

Section PINUS 

Subsection PONDEROSAE Loudon 

Pinus riogrande Axelrod 

Text figure 7C 

Pinus riogrande Axelrod, 1986 (part, typic), p. 624, fig. 68. 

Pinus sanjuanensis auct. non Axelrod 1986. Axelrod, 1987 (part), p. 95, pl. 13, 

fig. 1. 

Wolfe and Schorn (1989) designation: Pinus riogrande_. 

Description. -Ovulate cone shape narrow ovate 

(length to width ratio about 2:1 ); 12 em long, 4.5 em 

wide; apophysis in plane or facial view about 1.5 em wide 

and 0.8 em high, width to height ratio about 2:1; umbo 

dorsal, not markedly differentiated from apophysis, 

shape rhombic; mucro centro, straight or erect, about as 

long as height of umbo field. 

14 Spermatopsida of Oligocene Creede Flora, Colorado 

Discussion.-Included in this species is an 

impression of a complete ovulate cone, which is the 

holotype of Pinus riogrande, and a small fragment 

originally assigned to P. sanjuanensis. Both specimens 

share the features of (1) dorsal umbos that are weakly 

differentiated from the apophysis and (2) a mucro that is 

centroerect (situated on the medial keel and directed 

apically). In his description of the species, Axelrod (1986, 

p. 624) stated: "***umbo dorsal, centro-reflexed, short 

mucronate prickles 2-3 mm long, directed distally." This 

statement is confusing in the context of the terminology 

defined by Klaus (1980). 

Three additional ovulate cone fragments originally 

assigned to Pinus riogrande have well-defined umbos that 

protrude well above the plane of the apophysis and 

represent Pinus sp. 1. Three winged seeds previously 

referred toP. riogrande (Axelrod, 1987, pl. 12, figs. 2-4) 

have immature seeds and represent atypical seeds of an 

indetermined pine. 

Holotype.-UCMG 19704. 

Hypotype.- UCMP 7393. 

Pinus sp.l 

Text figure 7D 

Pinus riogrande Axelrod, 1986 (part, nontypic), p. 624, figs. 69, 70. Axelrod, 1987, 

text (part), p. 94, pl. 12, figs. 5, 6. 

Wolfe and Schorn (1989) designation: Pinus crossii. 

Description. -Ovulate cones incomplete, largest 

specimen 11 em long, 5.5 em wide; apophysis in plane or 

facial view 1.4 em wide, about 1.2 em high, width to 

height ratio about 1:1; umbo dorsal, distinct, projected 

above plane of apophysis; apophysis oblate, rhombic; 

mucro centro erect, about as long as height of upper field 

of umbo. 

Discussion.-Included here in Pinus sp. 1 are the 

apical portions of two ovulate cones originally assigned to 

P. riogrande. The incompleteness of these two specimens 

dictates against any serving as holotypic material for a 

named species. These two specimens apparently also 

represent a species of Ponderosae, although they are 

distinct from P. riogrande (compare fig. 7C to 7D). 

Specimens.- UCMP 7209, 7210. 

Pinus sp. 2 

Pinus macginitiei Axelrod, 1986 (part, nontypic), p. 621, fig. 56. Axelrod, 1987 

(part), p. 93, pl. 9, fig. 9. 

Wolfe and Schorn (1989) designation: Pinus crossii. 

Discussion. -A single Creede specimen of a disarticulated wing may represent an entity distinct from Pinus 

crossii and other described fossil taxa that are known 

from winged seeds. The wing is widest immediately distal 

to the seed area and was probably borne on a cone scale 

that was expanded near the middle of the scale and 

narrowed to either end. The wing of P. macginitiei from 
Florissant is not expanded, and presumably the bearing 

cone scale was of approximately uniform width; this 

difference between the Creede and Florissant winged 

seeds is considered sufficient to exclude the Creede 

specimen from P. macginitiei. 

This Creede winged seed has some similarities to 

seeds of subsection Ponderosae but is not sufficiently 

diagnostic to eliminate from consideration placement in 

other subsections. Possibly the Creede specimen 

represents the same entity as our Pinus sp. 1, but this 

suggestion is conjectural. 

Nomenclature.-This specimen (UCMG 19701) 

was cited in figure legends by Axelrod (1986, fig. 56; 

1987, pl. 9, fig. 9) as the "holotype" of Pinus macginitiei. 

Axelrod (1986, p. 621), however, designated as the 

primary type (erroneously called a "lectotype" rather 

than a holotype) UCMP 3776, which is a long (15 em), 

three-needled fascicle (persistent sheath) from Florissant 

originally attributed by MacGinitie (1953, pl. 20, fig. 1) to 

P. f/orissantii. In his validating description of P. macginitiei, Axelrod (1986, p. 621) first described the Florissant 

fascicle and then cited UCMG 19703 as a "hypotype" 

(properly, this is a paratype). Clearly the holotype of P. 

macginitiei is the Florissant fascicle, and the Creede 

winged seed is a paratype, which is here excluded from P. 

macginitiei. 

Specimen.- UCMG 19703. 

Subgenus lndetermlned 

Pinus Species lndetermined (atypical winged seeds and wings) 

PL'lte 1, figures 13, 14; plate 2, figure 10 

Picea coloradensis Axelrod, 1987 (part, nontypic), p. 84, pl. 8, figs. 9-14. 

Pseudotsuga glaucoides Axelrod, 1987 (part, nontypic), p. %, pl. 15, figs. 7, 8. 

Tsuga petranensis Axelrod, 1987 (part, nontypic), p. 98, pl. 7, fig.10 (figure legend 

uses the nomen nudum of Tsuga coloradensis). 

Picea lahontensis auct. non MacGinitie. Axelrod, 1987 (part), p. 85. 

Picea sonomensis auct. non Axelrod 1944. Axelrod, 1987 (part), p. 86, pl. 8, 

figs. 7, 8. 

Pinus alvordensis auct. non Axelrod 1944. Axelrod, 1986 (part), p. 627, figs. 77, 

82-84. Axelrod, 1987 (part), p. 89, pl. 9, figs. 5-7. 

Pinus coloradensis auct. non Knowlton. Axelrod, 1986 (part), p. 615, figs. 47, 49, 

50, 52. Axelrod, 1987, (part), p. 90, pl. 10, figs. 3, 6, 7. 

Pinus riogrande auct. non Axelrod 1986. Axelrod, 1987 (part), p. 94, pl. 12, 

figs. 2-4. 

Wolfe and Schorn (1989) designation: Pinus crossii. 

Discussion.-All the structures listed in the above 

synonymy are referable to Pinus. Attempts to match 

these to the typically developed winged seeds of various 

extant pinaceous genera results in unjustified illusions of 

similarities. As discussed previously, the wings and (or) 

winged seeds of Pinaceae (especially Pinus) are so highly 

variable that a single cone can produce a variety of 

atypical wings and (or) winged seeds that can be grossly 

matched with other species of pine. Such atypical winged 

seeds should be expected in the fossil record and should 

be evaluated relative to the more common, recognizably 

typical, winged seeds in the same fossil assemblage. Most 

atypical Creede winged seeds of Pinus probably 

represent the abundant P. crossii and were thus included 

in the Wolfe and Schorn (1989) revised census data 

under that taxon. 

Specimens.- UCMG 19706-19708; UCMP 

7228-7230, 7232, 7245 (counterpart of UCMP 7228) ), 

7278, 7296, 7299, 7309, 7318-7324 (7324=7319), 

7373-7376 (7373 = UCMG 19708), 7378, 7385, 7395, 

7397, 7430, 7433, 7434-7437, 7441-7444. Note that 

UCMP 7232 was figured by Axelrod (1986, fig. 77) as 

Pinus alvordensis and by Axelrod (1987, pl. 8, fig. 14) as 

Pice a coloradensis. 

Genera and (or) Species Indetermined (cone fragments) 

PL'lte 2, figures 5-9 

Picea co/oradensis Axelrod, 1987 (part, typic), p. 84, pl. 8, figs. 15, 16. 

Tsuga petranensis Axelrod, 1987 (part, typic), p. 98, pl. 7, figs. 7-9. 

Pseudotsuga glaucoides Axelrod, 1987 (part, nontypic), p. %, pl. 15, figs. 2, 5, 6. 

Picea lahontensis auct. non MacGinitie. Axelrod, 1987 (part), p. 85, pl. 8, fig. 1. 

Pinus coloradensis auct. non Knowlton. Axelrod, 1986 (part), pl. 615). Axelrod, 

1987 (part), p. 90. 

Pinus crossii auct. non Knowlton. Axelrod, 1987 (part), p. 91. 

Discussion.-All the cone remnants included in the 

above synonymy have been abraded biologically and (or) 

mechanically. Critical characters needed for generic 

determination are typically lacking (see previous 

discussion of abrasion states). A few remnants are 

probably sufficiently well preserved to indicate inclusion 

in Pinus, but specific assignment is impossible. 

The specimen that Axelrod (1987, pl. 7, fig. 9) 

considered to be an ovulate cone scale of Tsuga is a cone 

scale of Pinus. The specimen was figured with the apex 

pointing toward the bottom of the plate and is incorrectly 

retouched. The basal end of the cone scale (oriented 

toward the top of the plate) was eliminated by retouching 

and is frayed, as if the cone scale had been biologically 

removed from the parent ovulate cone. The structure 

that appeared from retouching to be seed cavities is the 

apophysis of the cone scale. 

Specimens.- UCMP 7199 (probably 

cone of Pinus), 7287-7294, 7314, 7356 

immature cone of Pinus), 7417-7424 

counterpart of 7417), 7438-7440. 

Family BERBERIDACEAE 

Genus BERBERIS Llnnaeus 

Berberis coloradensis Axelrod 

Plate 4, figures 1-7, 9 

Berberis coloradensis Axelrod, 1987, p. 110, pl. 22, figs. 1-3, 6, 7. 

Berberis riogrande Axelrod, 1987, p. 110, pl 21, figs. 8-11. 

immature 

(probably 

(7420 is 

Fendlera coloradensis Axelrod, 1987 (part, typic), p. 122, pl. 24, figs. 5-8. 

Philadelphus creedensis Axelrod, 1987, p. 124, pl. 24, figs. 1-4. 

Vaccinium creedensis Axelrod, 1987 (part, typic), p. 147, pl. 33, figs. 3, 4. 

Wolfe and Schorn (1989) designation: Berberis coloradensis. 

Emended description.-Leaves simple; shape 

symmetric, narrowly elliptic to oblanceolate; base 

Systematic Descriptions and Discussions 15 
symmetric, cuneate, apex acute to obtuse and mucronate; 

length 0.8-1.3 em, width 0.2-0.6 em; venation perfectly 

actinodromous to acrodromous; moderate midrib and 

two lateral primary veins; lateral primary veins typically 

extending half the distance to apex, where they form 

angularly brochidodromous loops with secondary veins 

about half the distance from midrib to margin; two to 

three pairs of medial secondary veins, originating at 

angles of 50°-70°, extending straight to form brochidodromous loops that are elongated parallel with midrib; 

loops abmedial to secondary loops festooned brochidodromous; if teeth present, an external secondary vein 

from lateral primary vein extends through medial part of 

tooth to end as spine; intercostal venation formed 

typically by a composite intersecondary vein that 

branches into randomly reticulate tertiary veins; fourthorder veins forming quadrangular to pentagonal, welldeveloped areoles; veinlets mostly once branching; 

margin has fimbria! vein, typically entire but can have one 

large spinose tooth on either or both sides; petiole 

absent. 

Discussion.-The five taxa synonymized here share 

several critical characters: (1) actinodromy, (2) only two 

lateral primary veins, (3) the lateral primary veins extend 

typically about half the distance to the apex where they 

are joined by strong veins that originate from the midrib, 

(4) festooned brochidodromy, (5) a fimbrial vein, (6) a 

mucronate (spinose) apex, and (7) absence of a petiole. 

Berberis coloradensis is typically entire-margined. The 

type suite of B. coloradensis comprises leaves that have a 

pair of spinose teeth and a mucronate (spinose) apex, but 

the type suite of B. riogrande was stated to differ in 

lacking teeth and in having a rounded rather than an 

apiculate (mucronate) apex {Axelrod, 1987, p. 111; 

"mucronate," which specifically refers to an extension of 

the midrib, is preferred to the more general "apiculate"). 

The photograph of the holotype of B. riogrande (Axelrod, 

1987, pl. 21, fig. 11) displays a small projection on the 

righthand side; cleaning revealed a large, spinose tooth 

(pl. 4, fig. 3). The apices of the holotype and most other 

primary types of B. riogrande are missing, but one paratype (pl. 4, fig. 2) has a mucronate apex. Similarly, 

Fend/era coloradensis and Philadelphus creedensis both 

have paratypes that, when cleaned, have single spinose 

teeth (pl. 4, figs. 4, 6) and mucronate apices (pl. 4, figs. 5, 

7). The holotype (pl. 4, fig. 9) and one paratype of 

Vaccinium creedensis, although somewhat broader than 

typical for B. creedensis and untoothed, have the seven 

critical characters listed above, including a mucronate 

apex; an additional paratype of V. creedensis is an 

undeterminable dicot lamina. 

Because Berberis coloradensis includes material 

referred to four different genera in three different 

families that represent subclasses Ranunculidae, Dilleniidae, and Rosidae, the proper supraspecific assignment 

16 Spermatopsida of Oligocene Creede Flora, Colorado 

of the species is open to question. Festooned brochidodromy is a primitive character in the dicotyledons 

(Hickey and Doyle, 1977) and is particularly common in 

Magnoliidae and Ranunculidae. Actinodromy associated 

with a fimbria} vein is particularly common in Ranunculidae (Lardizabalaceae, Menispermaceae, and Berberidaceae ), and spinose teeth occur throughout Berberidaceae. Of the three extant berberidaceous genera that 

are spinose, Epimedium has deeply cordate leaves that 

have more than one pair of lateral primary veins and 

numerous teeth, and Mahonia has leaves that are pinnately compound, which results in isolated leaflets that 

are typically asymmetric. The symmetry of the laminae of 

B. coloradensis indicates that the generic reference is 

valid, and the absence of a well-defined petiole is 

consistent with this; the laminae of extant Berberis are 

considered to be homologues of the apetiolulate terminal 

leaflet of Mahonia (Ahrendt, 1961). 

Extant foliage of Berberis varies in venational 

architecture from actinodromous (and, as in B. coloradensis, long and strong lateral primary veins occur) to 

predominantly pinnately veined; some spedes have an 

intermediate stage, in which lateral primary veins are 

greatly reduced in length and thickness and the medial 

part of the lamina is dominated by secondary veins that 

originate near (but not at) the laminar base. Because 

Mahonia appears, as do most ranunculid groups, to be 

primitively actinodromous, we infer that actinodromy is 

probably primitive in Berberis. 

Berberis coloradensis was compared only to B. 

candidula Schneid. (Axelrod, 1987), which is highly 

dissimilar from the fossil in being pinnately veined and 

having three or four teeth per laminar side. Berberis 

riogrande was compared to six extant species of Berberis 

(Axelrod, 1987), of which three belong to section Sinenses and two belong to section Asiaticae (Ahrendt, 

1961). Leaves of Sinenses represent either the 

intermediate or pinnately veined conditions, and leaves 

of Asiaticae are pinnately veined. The last supposedly 

comparable extant species has architecture of the 

intermediate stage, but section Angulosae, to which this 

species belongs, contains other species that have 

markedly actinodromous architecture (for example, B. 

diaphana Maxim. and B. sibirica Pall.). Laminae of 

subsections Diaphanae and Sibiricae, however, typically 

have four or more teeth (Ahrendt, 1961). 

Some members of section Polyanthae (for example, 

Berberis wilsoniae Hemsl.) have major venational and 

marginal architecture similar to B. coloradensis, but the 

secondary and tertiary veins are thick and approximately 

the same width as the lateral primary veins (the 

"reticulate venation" of Ahrendt, 1961). In B. coloradensis, the tertiary veins are markedly thinner than lower 

order veins, a condition found in other Ranunculidae and 

thus probably plesiomorphic. 
Relative to minor as well as major venational and 

marginal architecture, Berberis coloradensis is probably 

most similar to leaves of subsection Eufranchetianae of 

section Franchetianae. Berberis amoena Dunn, for 

example, has venation markedly like the Creede species; 

although B. amoena is entire-margined, other related 

species rarely have one to four teeth per side (Ahrendt, 

1961). We question, however, whether B. coloradensis 

should be assigned to Franchetianae: (1) marked actinodromy and differentiation of major and minor venation 

are interpreted as plesiomorphies rather than as synapomorphies, and (2) few to no teeth in Berberis appear to 

be correlative with reduction in laminar size in response 

to a cold environment. Perhaps significant is that floral 

and fruit characters that could be assumed to be plesiomorphies in Berberis (for example, "subracemose" inflorescences [Ahrendt, 1961] and several or more ovules) 

are also found in sections Franchetianae and Angulosae. 

The inferred plesiomorphies of B. coloradensis foliage 

make problematic assignment of this species to any 

extant section of the genus. 

Holotype.-UCMP 7527. 

Paratypes.-UCMP 7525, 7526, 7528, 7529. 

Hypotypes.-VCMP 7530, 7534,7653, 7655-7666, 

7745-7753, 7849, 7850. 

Genus MAHONIA Nuttall 

Mahonia aceroides (Knowlton) Schorn & Wolfe, n. comb. 

Plate 4, figures 8, 10, 11; text figure 8 

Sterculia aceroides Knowlton, 1923, p. 191, pl. 43, fig. 12. 

Mahonia creedensis Axelrod nom. nud., 1956, p. 293. Axelrod, 1987, p.111, pl. 23, 

figs. 1-16. 

Odostemon hakeaefolia (Lesquereux) Knowlton, 1923 (part, nontypic), p.190, pl. 

43, figs. 1-6. 

Odostemon marginata (Lesquereux) Knowlton, 1923 (part, nontypic), p. 188, pl. 

43, figs. 7-10. 

Mahonia obliqua auct. non MacGinitie. Axelrod, 1987, p.l14, pl. 21, figs. 12-15. 

Wolfe and Schorn (1989) designation: Mahonia sp. 

Description.-Leaves odd-pinnate with at least 

seven leaflets; leaflet shape ovate to narrowly oblong; 

base asymmetric, obtuse to broadly rounded, apex acute 

and spinose; length 1.1-4.0 em, width 1.1-2.2 em; some 

terminal leaflets have basal lateral lobes; venation 

perfectly actinodromous, three to five primary veins; 

midrib moderate, typically slightly to markedly curving; 

most apical pair of lateral primary veins extending onethird or more distance to apex, typically forming a 

brochidodromous loop with a medial secondary vein and 

the loop elongated parallel to midrib, rarely craspedodromous and entering teeth medially; most basal pair of 

lateral primary veins, if present, typically craspedodromous into most basal teeth; veins entering teeth apical to 

most basal teeth typically originate as either external 

veins from primary-vein loops or as medial secondary 

veins or as external veins from secondary-vein loops; 

intersecondary veins composite, common, some 

extending to brochidodromous loops; third-order venation randomly reticulate; fourth-order veins forming 

well-developed, typically quadrangular areoles; veinlets 

mostly unbranched or once branched; fimbrial vein; two 

to five large, spinose, C-2 to typically C-3 teeth on each 

side of lamina, irregularly spaced, typically in basal 

two-thirds of lamina; sinuses markedly arcuate; apetiolulate. 

Discussion. -Odd-pinnate leaves that have actinodromous terminal and lateral leaflets are known only in 

Mahonia. Presence of a few spinose teeth and a fimbria} 

vein further indicate validity of the generic assignment 

made by Axelrod (1987). 

Mahonia obliqua has an exceedingly narrow 

lamina, as pointed out by MacGinitie (1953); such laminae were validly determined by Becker (1962) in the 

Ruby flora. The leaves assigned by Axelrod (1987) toM. 

obliqua, however, are either too broad (his pl. 21, figs.l2, 

13), or are folded (his pl. 21, fig.l4), or are a fragment of 

a larger lamina (his pl. 21, fig. 15). These Creede 

specimens probably represent M. aceroides. 

Nomenclature.-Knowlton's type of Sterculia aceroides (pl. 4, figs. 8a, lOa) represents the basal part of an 

actinodromous lamina that appears to have "teeth" 

atypically large for the Creede Mahonia. The "teeth," 

however, are similar in size and shape to basal lateral 

lobes that can be found on terminal leaflets of extant 

species such as the western American M. neNosa Pursh; 

in other characters (for example, areolar venation; fig. 8), 

the type of S. aceroides can be matched by other Creede 

specimens of Mahonia. The specimen of S. aceroides has 

never been completely cleaned because to do so would 

require partial destruction of an adjacent specimen (pl. 4, 

figs. 8b, lOb), which is one of Knowlton's (1923) 

illustrated specimens of Odostemon hakeaefolia (a typical 

lateralleaJflet of the Creede Mahonia). Because only one 

species of Mahonia is present in the Creede flora, 

retention of the epithet aceroides should not lead to error 

or confusion; the abundant material of Mahonia 

available from Creede allows full characterization of the 

foliage of .M. aceroides .. 

Although Mahonia creedensis was published by 

Axelrod (Jl956) as a "new name," the lack of a previously 

Figure 8. Drawing showing areolar venation of Mahonia. A, 

M. aceroicfes (Knowlt.) Schorn & Wolfe, holotype USNM 

36537 (sarne specimen as pl. 4, figs. Sa, 10a). 8, M. aceroides (Knowlt.) Schorn & Wolfe, hypotype USNM 36531 

(same specimen as pl. 4, figs, 8b, 1 Ob). 

Systematic Descriptions and Discussions 17 
published description and (or) diagnosis clearly, under 

the ICBN, made this a new species; for a new species, a 

holotype is selected, not a lectotype. A description and 

(or) diagnosis of M. creedensis was not published by 

Axelrod (1956, 1987), and therefore this binomial must 

be rejected as illegitimate. Indeed, the inclusion of Sterculia aceroides in Axelrod's (1956) synonymy of M. 

creedensis, although now rejected by him (Axelrod, 

1987), made M creedensis a superfluous name at the 

time of publication. We note also that, in continuing 

contravention of the ICBN, Axelrod (1987, p. 113) 

proposes another illegitimate binomial in Mahonia: M. 

harneyensis Axelr. is unaccompanied by either a 

description or a reference to a previously published 

description. 

Holotype.-USNM 36537. 

Hypotypes.-USNM 36526-36535; UCMP 7537-

7571; UCMG 19749. 

Family SALICACEAE 

Genus POPULUS Linnaeus 

Populus larsenil (Knowlton) Schorn & Wolfe, n. comb. 

PL'lte 5, figures 1, 2, 5; text figure 9 

Alnus? /arseni Knowlton, 1923, p. 188, pl. 42, fig. 3. 

Populus creedensis Axelrod, 1987, p. 103, pl. 18, figs. 1-4; pl. 19, figs. 1-4; pl. 21, 

fig. 1. 

Populus lesquereuxi auct. non Cockerell. Knowlton, 1923, p. 187, pl. 44, fig. 1. 

Populus cedmsensis auct. non Wolfe. Axelrod, 1987, p. 101, pl. 19, fig. 5; pl. 20, 

fig. 3. 

Populus pliotremuloides auct. non Axelrod, 1937. Axelrod, 1987, p. 104, pl. 20, 

fig. 1. 

Betula smithiana (Axelrod) Axelrod, 1987 (part, nontypic), p. 106, pl. 21, 

figs. 7a, 7b. 

Wolfe and Schorn (1989) designation: Populus sp. 

Description. -Leaves simple; shape symmetric, 

ovate to narrowly ovate; base symmetric, obtuse to 

broadly rounded, apex acute to attenuate; length 4.0-

> 10 em, width 2.5-5.0 em; venation pinnate, midrib 

Figure 9. Drawing showing venation of Populus larsenii 

(Knowlt.) Schorn & Wolfe. USN M 36519 (holotype of Alnus? 

larseni Knowlt.). 

18 Spermatopsida of Oligocene Creede Flora, Colorado 

moderate; typically with two pairs of secondary veins 

originating decurrently close to base and extending about 

one-fourth to one-half the distance to apex; four to six 

pairs of more apical secondary veins, originating slightly 

decurrently at angles of 30°-50°, broadly convex, weakly 

brochidodromous to eucamptodromous; intersecondary 

veins simple, common, some extending more than half 

the distance to margin; tertiary veins A-R to A-0, 

branching, somewhat sinuous, percurrent; fourth-order 

veins at variable angles to tertiary veins, forming large, 

imperfect, irregularly polygonal areoles; veinlets dendritically branching; weak external secondary veins, 

especially originating from two most basal pairs of 

secondary veins, typically looping; teeth entered by weak 

veins that originate either as external secondary veins or 

typically as branches from loops of external secondary 

veins; teeth typically at least three to four per secondary 

vein (more near base), C-1, salicoid; sinuses angular; 

petiole 1.3-4.1 em long. 

Discussion.-The Creede Populus and many extant 

species of the genus typically have two pairs of strong, 

decurrent secondary veins that originate close to the base 

of the lamina; presumably this architecture is derived 

from the actinodromous condition found in some Populus and the related Flacourtiaceae. Well-preserved 

specimens of the Creede Populus have salicoid teeth 

(Hickey and Wolfe, 1975), and many specimens also have 

acropetiolar glands that commonly occur in extant Populus. 

Within a given species of extant Populus, leaves can 

vary in size, shape, and spacing and size of teeth along a 

given shoot (S.L. Wing, oral commun., February 1989). 

Leaves of young plants can differ even more markedly 

from mature leaves of mature plants. Some suites of 

leaves can be interpreted as representing mature leaves 

of mature plants, as in the instance of the suite described 

and illustrated as P. creedensis, but attributing single 

specimens in the same assemblage to additional species 

of the genus is very questionable. The Creede specimens 

attributed to P. cedlUsensis and P. pliotremuloides are 

considered here probably to be extreme variants of P. 

larsenii. 

The specimen assigned to Populus cedlUsensis 

differs in several respects from that species, which is 

known from a large suite of specimens (Wolfe, 1964; 

Schorn, unpublished data): (1) P. cedlUsensis is truly 

actinodromous rather than having one or two pairs of 

strong suprabasal secondary veins, (2) teeth in P. 

cedlUsensis are very reduced and have a basal side 10 to 

20 times as long as the apical side, rather than distinct 

teeth that have a basal side only 2 to 3 times as long as the 

apical side, (3) teeth in P. cedlUsensis are only about one 

or two per secondary vein, rather than two or three, and 

(4) the lateral primary veins in P. cedlUsensis typically 
extend about half the distance to the apex in contrast to 

suprabasal secondary veins that extend about one-fourth 

to one-third the distance to the apex. 

Axelrod (1987) attributed two fragmentary 

specimens to Betula smithiana (Axelr.) Axelr. Both 

specimens have brochidodromous rather than craspedodromous secondary veins (and hence cannot represent 

Betula) and have weak external veins that enter small 

narrowly rounded, closely spaced teeth. A swelling at the 

top of the petiole of one specimen (UCMP 7512) 

suggests the acropetiolar glands commonly present in 

Populus. 

The leaves of Populus larsenii resemble both the 

extant P. trichocarpa Torr. & Gray and P. balsamifera L.; 

however, as noted by Axelrod (1987), the associated 

two-valved capsules indicate a relationship to P. balsamifera rather than to P. trichocarpa, which has three-valved 

capsules. 

Nomenclature.-The type of Alnus? larseni has 

strong, decurrent suprabasal secondary veins, ovate 

shape, and numerous salicoid teeth; venation is 

preserved to the fourth order (fig. 8) and conforms to 

venation in Populus creedensis. No doubt can exist about 

the generic affinities of this specimen, and retention of 

the epithet larsenii will not lead to error and (or) 

confusion. 

Holotype.-VSNM 36519. 

Hypotypes.-VCMP 7476--7494, 7512, 7513; 

UCMG 34060, 34061. 

Genus SALIX Linnaeus 

Salix sp. 

Salix venosiuscula auct. non H.V. Smith. Axelrod, 1987, p. 106, pl. 20, figs. 7, 8. 

Wolfe and Schorn (1989) designation: Salix sp. 

Discussion.-Salix venosiuscula was founded 

(Smith, 1941) on leaves that have (1) an elliptic to slightly 

obovate shape, (2) an acute apex, (3) a length to width 

ratio of about four to one, ( 4) about 15 pairs of secondary 

veins, ( 5) secondary veins that originate at angles of 

about 50° from the base to the apex, and (6) a closely and 

uniformly serrate margin. In contrast, the Creede 

specimens assigned to this species have (1) a markedly 

ovate shape, (2) a very attenuated apex, (3) a length to 

width ratio of more than five to one (the laminae are 

incomplete), ( 4) more than 25 pairs of secondary veins, 

(5) secondary veins that originate at angles of about 

40°-60° basally and about 80° apically, and (6) a sparsely 

and irregularly serrate margin. Although the Creede 

leaves cannot represent S. venosiuscula, their 

preservation is too poor for valid specific determination 

or to suggest affinities to extant species. 

Specimens.- UCMP 7507-7511. 

Family PHH.ADELPHACEAE 

Genus JAMESIA Torrey & Gray 

Jamesla caplanii Axelrod 

Plate 5, figure:; 3, 4 

lamesia caplanii Axelrod, 1987, p. 123, pl. 30, figs. 12-16. 

Wolfe and Schorn (1989) designation: lamesia caplanii. 

Description. -Leaves simple; apparently estipulate; 

shape symmetric, elliptic to typically slightly ovate; base 

symmetric:, obtuse to rounded, some slightly cordate; 

apex acute: to slightly acuminate; length 1.4-4.2 em, width 

0.9-2.4 em; venation pinnate; midrib moderate; five to 

seven pairs of secondary veins, thin, more basal pairs 

decurrent, originating at angles of as much as 85° near 

base and decreasing to 20° near apex, broadly convex, 

some brochidodromous but most craspedodromous, 

entering tt::eth medially; tertiary veins A-R toR-R, widely 

spaced, reticulate; fourth-order veins forming imperfect, 

irregularly polygonal areoles; veinlets unbranching or 

typically once branching; zero to two external secondary 

veins originating at acute angles, broadly convex, craspedodromous; margin with uniformly spaced, large teeth 

that are one to two per secondary vein, D-1; teeth have 

lateral accessory veins convergent on an apical foramen; 

sinuses angular; petiole 0.3-1.5 em long, with five 

discrete vascular strands. 

Discussion-Markedly rosoid teeth (Hickey and 

Wolfe, 1975), as occur in these Creede leaves, are 

typically associated with actinodromy (see later 

discussion of Ribes ). Some pinnately veined, simple 

leaved Rosaceae can have somewhat weakly developed 

rosoid teeth, but in those taxa intersecondary veins are 

common and the tertiary veins are typically not oriented 

almost orthogonally to the midrib as in Jamesia caplanii. 

Orthogonal orientation of the tertiary veins relative to 

the midrib, absence of intersecondary veins, a simple leaf, 

and eucamptodromy are characteristic of many advanced 

Rosidae (.for example, Cornales, Myrtales, Rhamnales, 

Celastrales, and Santalales) and are found in some basal 

rosid groups. These basal groups include Iteaceae and 

Hydrangeaceae, and this architecture is therefore termed 

iteid. The iteid architecture is typically associated with 

eucamptodromy, and if the margin has teeth, the teeth 

are small. Only in Jamesia is iteid intercostal venation 

combined with large, rosoid teeth and craspedodromy of 

the secondary veins. 

Most specimens of Jamesia caplanii are markedly 

similar to the extant J. americana Torr. & Gray in all 

aspects of laminar vasculature and morphology of the 

teeth; however, even small leaves of J. caplanii (pl. 5, fig. 

4) have five discrete vascular bundles in the petiole as 

opposed to three or four in the extant taxon, and the 

extant taxon typically has a shorter petiole. Laminae and 

petioles of J. americana are densely pubescent; although 

some specimens of J. caplanii are preserved in a fineSystematic Descriptions and Discussions 19 
Table 2. Differentiation of Creede species of Ribes 

Character Ribes /acustroides 

Orbiculate 

5 

(Shallow-)deep 

R. robinsonii 

Widely ovate 

5 

Shallow 

R. obovatum 

Obovate 

0 

Laminar shape 

Number of lobes 

Lobal dissection 

Lobal apex 

Lobal shape 

Base shape 

Stipules 

Narrowly to broadly obtuse 

Triangular to trapezoidal 

Deeply cordate 

Narrowly obtuse 

Triangular 

Shallowly cordate 

Small 

None 

None 

None 

Acute to obtuse 

Large ? 

grained matrix, no evidence of dense pubescence is 

preserved. More significantly, some specimens of l 

caplanii (for example, pl. 5, fig. 3) have small, closely 

spaced teeth and some secondary veins that are almost 

eucamptodromous. This morphology is more like that 

found in Iteaceae and Hydrangeaceae and suggests 

retention of some primitive characters in foliage of J. 

caplanii. 

Excluded from Jamesia caplanii is Becker's (1962, 

pl. 18, fig. 3) Ruby specimen originally referred to Ulmus 

paucidentata H. V. Sm. and included by Axelrod (1987) 

as a paratype of the Creede Jamesia. The Ruby specimen 

has markedly arcuate dental sinuses and cannot 

represent Jamesia, which has markedly angular dental 

sinuses. 

Holotype.-UCMP 7641. 

Paratypes .-U CMP 7642-7651. 

Hypotypes.-UCMG 34062,34063. 

Family GROSSULARIACEAE 

Genus RIBES Linnaeus 

Discussion-One of the most common 

dicotyledonous leaf types in the Creede collections is 

leaves that are actinodromous, palmately lobed, and 

serrate or crenate. This leaf type accounts for 25 percent 

of Axelrod's (1987) species of Creede dicotyledons and 

was split into 13 species representing 5 genera and 4 

families: Ranunculus (1 species; Ranunculaceae ), Ribes 

(8 species; Grossulariaceae ), Physocarpus (2 species; 

Rosaceae), Rubus (1 species, Rosaceae), and Acer (1 

species; Aceraceae). A number of morphologic criteria 

indicate that most of these generic and familial 

assignments must be erroneous and that far fewer 

species are represented. 

Ranunculaceae have chloranthoid teeth (Hickey 

and Wolfe, 1975), which are lacking in the single 

specimen of Ranunculus creedensis Axelr. Further, 

Ranunculaceae are estipulate (Airy Shaw, 1985); 

cleaning of the specimen of R creedensis revealed a 

stipulate petiole 1.7 em long. Similarly, the holotype of 

Acer riogrande as illustrated by Axelrod (1987) appears 

to be stipulate, whereas all extant Acer are estipulate 

(Airy Shaw, 1985). Indeed, where the base of the petiole 

is preserved on the Creede actinodromous, lobed leaves, 

20 Spermatopsida of Oligocene Creede Flora, Colorado 

stipules are present. Aside from the two specimens just 

mentioned, also stipulate are specimens of Ribes dissect a, 

R lacustroides, R. stevenii, R webbii, R. riogrande, R 

birdseyii, and Physocarpus petiolaris. Five of Axelrod's 

species of Ribes were considered to be closely related to 

extant species that fall into subgenus Ribes, a taxon that 

typically is estipulate. Indeed, stipules are uncommon in 

extant Grossulariaceae and apparently occur only in 

some species of subgenus Grossularia of Ribes. The other 

two of Axelrod's actinodromous, lobed species in the 

Creede flora (R. wasonii and P. triloba) are represented 

by single specimens that have incomplete petioles. 

The rosoid tooth (Hickey and Wolfe, 1975) occurs 

in three families associated with actinodromous, lobed 

leaves: Grossulariaceae, Rosaceae, and Vitidaceae. All 

the Creede lobed, actinodromous leaves have rosoid 

teeth and thus must represent one or more of these three 

families. The Creede leaves, moreover, have crenations, 

and thus Vitidaceae, which typically have broad A-1, 

D-1, or D-4 serrations, are eliminated from 

consideration. Further, Vitidaceae typically do not have 

subsidiary teeth, whereas all the Creede leaves under 

consideration here have subsidiary teeth. 

In Grossulariaceae (as well as Vitidaceae ), the 

rosoid tooth is what could be termed classic, with two 

strong lateral subsidiary veins approaching and ending at 

the apical foramen (see pl. 9, fig. 1). In Rosaceae, the 

lateral subsidiary veins tend to be weak and typically thin 

conspicuously before reaching the foramen (see pl. 9, fig. 

8); in many rosaceous taxa the veins fail to reach the 

foramen and (or) a foramen is absent. In the better 

preserved Creede specimens, most have the classic 

rosoid tooth and thus represent Grossulariaceae. Ranunculus creedensis has weakly developed rosoid teeth, 

however, and is later transferred to Potentilla in 

Rosaceae. Only three, rather than eight, species of Ribes 

are considered to be represented in the Creede flora 

(table 2), and two of these were assigned by Axelrod 

(1987) to Rosaceae. 

Ribes lacustroides Axelrod 

PL'lte 5, figures 6, 7, 10, 11; plate 6, figures 1-17 

Ribes lacustroides Axelrod, 1987, p. 118, pl 24, figs. 16--18. 

Ribes birdseyii Axelrod, 1987, p. 115, pl. 32, figs. 7-9. 

Ribes creedensis Axelrod, 1987, p. 116, pl. 28, figs. 12, 13. 

Ribes dissecta Axelrod, 1987, p. 117, pl. 24, figs. 14, 15; pl. 25, fig. 10. 
Ribes riogrande Axelrod, 1987, p. 119, pl. 25, figs. 11-13. 

Ribes stevenii Axelrod, 1987, p. 120, pl. 25, figs. 1--6. 

Ribes wasonii Axelrod, 1987, p. 121, pl. 25, fig. 9. 

Rubus riogrande Axelrod, 1987, p. 137, pl. 31, fig. 1. 

Acer riogrande Axelrod, 1987, p. 141, pl. 32, fig. 10. 

Ribes protomelaenum auct. non Cockerell. Knowlton, 1923, p. 188, pl. 42, figs. 

5-9. 

Vitis florissantella auct. non Cockerell. Knowlton, 1923, p. 189, pl. 42, fig. 4. 

Ribes webbii auct. non Wolfe. Axelrod, 1987, p. 121, pl. 25, figs. 7, 8. 

Wolfe and Schorn (1989) designation: Ribes lacustroides. 

Emended description.-Leaves simple; stipulate; 

shape symmetric, orbiculate; base symmetric, shallowly 

to typically deeply cordate, apex narrowly to broadly 

obtuse; length 1.0-3.5 em, width 1.2-8.0 em; five lobed, 

more apical pair of lobes moderately to deeply incised 

and apices of. these typically at widest part of lamina; 

m.ore basal pau of lobes shallowly incised, typically only 

shghtly larger than compound teeth of suprajacent lobes; 

lateral lobes broadly triangular to somewhat convex sided 

to trapezoidal, apices narrowly to broadly obtuse; apical 

lobe ovate to broadly ovate; venation perfectly actinodromous but most basal pair of lateral primary veins can be 

pa!inactinodromous; midrib moderate; five primary 

v~ms; ~ateral primary veins straight to shallowly concave, 

dtvergmg from medial secondary veins; three to four 

(rarely as many as six) secondary veins originating from 

basal sides of lateral primary veins and from both sides of 

mi?rib; .one to five secondary veins originating from 

aptc~l s.Ides of lateral primary veins; secondary veins 

ongmatmg at angles of 20°-30°, straight to slightly 

convex, entering teeth medially, craspedodromous; lobal 

sinal braces formed by external secondary branches 

originating from either suprajacent or subjacent 

secondary vein; tertiary veins A-A, reticulate; fourthorder veins forming imperfect, irregularly polygonal areales; veinlets dendritically branching; medial lobe and 

basal sides of most apical pair of lateral lobes have three 

to four (rarely as many as six) crenate teeth; apical sides 

of most apical pair of lateral lobes have one to three 

crenate teeth (rarely as many as five); apical sides of most 

basal pair of lateral lobes typically have one crenate tooth 

(rarely as many as three teeth); large teeth may have one 

to two small crenate subsidiary teeth; large teeth 

markedly rosoid; sinuses angular; petiole 0.8-3.5 em 

long. 

Discussion. -Some small leaves of Ribes lacustroides have a superficial resemblance to leaves of R. webbii 

and were so referred by Axelrod (1987). Leaves of R 

webbii are ovate rather than elliptic, however, and the 

widest part of the lamina is on a line drawn between the 

apices of the more basal pair of lobes. The medial lobe of 

R. webbii is very shallowly incised, and the lobe narrows 

from the subjacent lobal sinuses; in these small leaves of 

R. lacustroides, the medial lobe is broadly elliptic, and the 

lobe first slightly widens apically from the subjacent lobal 

sinuses. 

As conceptualized here, Ribes lacustroides is highly 

variable relative to degree of laminar dissection between 

lobes and degree of rounding of lobes and teeth. 

Specimens that we assign to this taxon were assigned by 

Axelrod (1987) to 10 different species; specific 

differentiation was, in part, based on characters such as 

degree of laminar dissection, whether the major teeth 

have subsidiary teeth, and relative rounding of the lobes 

and teeth. However, the degree of laminar dissection, for 

example, represents a normal, if somewhat skewed, 

distribution for a single species (fig. 10). As defined by 

Axelrod (1987), R.lacustroides was, in part characterized 

by absence of subsidiary teeth, but examination of the 

holotype (pl. 6, fig. 2) indicates that some subsidiary teeth 

are present. 

Generally, the greater the dissection between lobes, 

the sharper the lobes and teeth appear; that is, as the 

sinuses adjacent to a lobe or tooth become more incised 

the lobe or tooth is more elongated and hence appear~ sharper. On plate 6, a series of specimens form a 

transition from the deeply dissected and "sharply" lobed 

and toothed R dissecta to the shallowly lobed and bluntly 

lobed and toothed R webbii (using Axelrod's, 1987 

designations); even if teeth appear sharp, examinatio~ shows them to be narrowly rounded. The forms referred 

toR creedensis (pl. 5, figs. 6, 7) represent a morphotype 

more extreme in deep dissection than R. dissecta. 

Between these extreme morpho types called R creedensis 

and R webbii are specimens referred to R dissect~ R. 

lacustroides, R stevenii, R. birdseyii, and R. riogrande; the 

single specimen that forms the basis for R. wasonii is 

aberrant and was damaged (probably by insects) during 

growth. The central morphotype in this transitional series 

is R. stevenii, which also comprises th~ most abundant 

morpho type. 

Some of the most aberrant leaves of Ribes lacustroides represent the morphotype called Rubus riogrande 

(pl. 5, figs. 10, 11); this is best represented (as are the 

30 

CJ) 

z 

UJ 

~ 

u 20 UJ 

a.. 

CJ) 

u. 

0 

f--

z 10 UJ 

u 

a: 

UJ 

a.. 

b=~!Z1itii 

0 0.1 0 .2 0.3 0.4 0.5 0.6 0.7 0.8 0 .9 1.0 

DEGREE OF DISSECTION (AlB) 

Figure 10. Histogram of laminar dissection in Ribes /acustroides Axelr. Horizontal axis is distance from sinus to top of 

pet~ole (A) divided by distance from lobe apex to top of 

petiole (B). The more deeply dissected specimens are to the 

left. Based on 181 specimens. 

Systematic Descriptions and Discussions 21 
more deeply dissected morphotypes of Ribes creedensis) 

at Axelrod's (1987) localities 571-A and 571-B, where 

Ribes leaves are most abundant and the greatest variation 

might be expected. The morphological intergradations 

between most of Axelrod's (1987) species of Creede 

Ribes indicate that one species is represented. As 

discussed earlier (p. 2), a variable taxon such as R. 

lacustroides (as we conceptualize it) could ultimately 

have given rise by canaliz-ation to two or more of the 

extant species of Ribes to which Axelrod 

(1987) compared different subsets of specimens of R. 

lacustroides. 

The single lamina on which Acer riogrande was 

based has not been deposited in the University of 

California Museum of Paleontology and thus has not 

been examined by us. The illustration (Axelrod, 1987, pl. 

32, fig. 10) appears to indicate a stipulate leaf, which 

would immediately eliminate Aceraceae. Additionally, 

one of the most apical lateral primary veins markedly 

diverges from the medial secondary veins (as in Grossulariaceae and Rosaceae), whereas inAcer the most apical 

lateral primary veins parallel (or slightly converge with) 

the medial secondary veins (Wolfe and Tanai, 1987). The 

specimen thus cannot represent Acer and probably 

represents a lamina of Ribes lacustroides, in which the 

margin is poorly preserved. 

Nomenclature.-We would have preferred to use 

the epithet stevenii for this taxon; of all the items included 

in the above synonymy, this is represented by the largest 

and best preserved suite of primary types. Axelrod (1987, 

p. 120), however, cited as the "lectotype" for Ribes 

stevenii the same specimen number as the "lectotype" for 

the previously instituted R. riogrande. Thus, being 

typified by the same specimen, R. stevenii immediately 

became a junior synonym of R. riogrande. The suites of 

primary types of R. birdseyii, R. creedensis, and R. dissecta 

are small, and we suggest selection under the ICBN of 

the epithet lacustroides. 

Holotype.-UCMP 7594. 

Paratypes.-UCMP 7585-7593, 7595, 7596, 7598, 

7601, 7606, 7609, 7620, 8599. 

Hypotypes.-UCMP 7577-7629, 7631-7638, 

7820A, B, 7836-7838, 7860; USNM 36523-36525; 

UCMG 27388, 27389, 27391, 34065, 34066. 

Ribes robinsonil Schorn & Wolfe, n. name 

Plate 5, figures 8, 9 

Physocarpus petiolaris Axelrod, 1987, p. 133, pl. 31, fig. 10. 

Physocarpus triloba Axelrod, 1987, p. 134, pl. 31, fig. 9. 

Wolfe and Schorn (1989) designation: Ribes sp. 1. 

Description. -Leaves simple; stipulate; shape 

symmetric, widely ovate; shallowly five lobed; most basal 

pair of lobes about same size as compound teeth of 

22 Spermatopslda of Oligocene Creede Flora, Colorado 

suprajacent lobes; base symmetric, shallowly cordate, 

apex narrowly obtuse; length 3.2-3.7 em, width 2.4-2.8 

em; venation perfectly actinodromous; midrib moderate; 

five primary veins; lateral primary veins straight to 

slightly concave or convex, diverging from medial 

secondary veins; three to four secondary veins originating 

from basal sides of lateral primary veins and from both 

sides of midrib; two to three secondary veins originating 

from apical sides of lateral primary veins; secondary veins 

originating at angles of 30°-40°, straight to slightly 

concave or typically convex, entering teeth medially, 

craspedodromous; lobal sinal braces formed by 

secondary veins originating from subjacent primary vein; 

tertiary veins R-R, branching; higher order venation not 

preserved; medial lobe and basal sides of lateral lobes 

have three to four large crenations; apical sides of most 

apical pair of lateral lobes have one to two crenations; 

apical sides of most basal lateral lobes typically have no 

teeth; large teeth have no subsidiary teeth on apical sides 

and zero to one on basal sides; sinuses angular; petiole 

2.0-2.9 em long. 

Discussion.-In leaves of Physocarpus (as well as in 

many Ribes ), the most basal lobes are entered by primary 

veins that originate as external veins from the two lateral 

primary veins (that is, the lamina is partially palinactinodromous ), whereas in Ribes robinsonii the lamina is 

perfectly actinodromous. Foliage that has characters 

similar to those listed for R. robinsonii (table 2) occurs in 

extant species such as R. amarum McClat. The petiole of 

one specimen of R. robinsonii (pl. 5, fig. 9) is densely 

covered with flat-topped hairs; the tops presumably bore 

glands. 

Nomenclature.-The epithets petio/aris and triloba 

(properly petiolare and trilobum in Physocarpus) have 

both been previously proposed in Ribes, and thus a new 

epithet is necessary. Peter Robinson has been of great 

assistance to this study by allowing ready access to the 

University of Colorado's Creede collections. 

Holotype.-UCMP 7817. 

Hypotypes.-UCMP 7630; UCMG 34064 

(counterpart USNM 422591). 

Ribes obovatum Schorn & Wolfe, n. sp. 

Plate 9, figures 1, 2 

Holodiscus hameyensis auct. non Chaney & Axelrod. Axelrod, 1987 (part), p. 130. 

Wolfe and Schorn (1989) designation: Ribes sp. 2. 

Description.-Leaves simple; shape symmetric, 

narrowly obovate; base symmetric, acute to obtuse; apex 

narrowly obtuse; length 2.5 em, width 1.5 em; venation 

perfectly actinodromous to acrodromous; midrib 

moderate; five primary veins; primary veins extending 

straight, either entering teeth medially and craspedodromous or branching and brochidodromous; all lateral 

primary veins extending at acute angles to the midrib at 
least one-half the distance to apex; one pair of medial 

secondary veins the same thickness as lateral primary 

veins originating at an angle of 15°, straight, entering 

teeth medially, craspedodromous; some composite intersecondary veins, extending about one-fourth the distance 

to margin; tertiary veins A-R, reticulate; fourth-order 

veins forming well-developed, quadrangular to 

pentagonal areoles; veinlets branching one to four times; 

dental sinuses braced by external veins that originate 

acutely from major suprajacent or subjacent veins; three 

large A-1 teeth on each side of lamina, crenate to slightly 

serrate; craspedodromous veins flanked by conspicuous 

veins that converge on foramen at apex of tooth; a single 

subsidiary tooth on basal side of most basal tooth on one 

specimen; sinuses angular; petiole incomplete, > 0.2 em 

long. 

Discussion.-The holotype and one paratype of 

Ribes obovatum were two of Axelrod's (1987) unfigured 

hom eo types of Holodiscus hameyensis. Ribes obovatum is 

clearly actinodromous in contrast to the pinnate venation 

of H o/odiscus. 

Ribes obovatum is unique in fossil Ribes in having a 

totally unlobed lamina. The leaves of the extant R. 

speciosum Pursh are typically broad and very shallowly 

three lobed, but some laminae have lobes the same size 

as teeth. Leaves of R speciosum can also have a single 

pair of medial secondary veins and three large teeth 

(counting the reduced lobes as teeth) per laminar side. If 

R speciosum and R obovatum are closely related, then R. 

obovatum would have to be considered as having the 

more reduced (specialized) lamina. 

Holotype.-UCMP 7788. 

Paratypes.-UCMP 7790; UCMG 34067; USNM 

422589. 

Family ROSACEAE 

Discussion.-Foliage of a large family such as 

Rosaceae is impossible to characterize, although some 

general characters apply to many, if not most, rosaceous 

taxa. Typically, teeth are a weakly developed rosoid type, 

in which the vein enters the teeth medially, but the lateral 

accessory veins are thin and may not reach the dental 

apex; a foramen at the dental apex is typically absent, 

although a small, dark conical gland may be present 

(especially in Pyroideae ). Small, dark conical glands 

(nectaries) occur on teeth near the base of the lamina in 

Prunoideae, as well as some laminar glands near the 

midrib in some Prunus. Although the cunonioid tooth 

(Hickey and Wolfe, 1975) is absent, some Rosaceae have 

the craspedodromous vein somewhat displaced toward 

the apical side, and some also have the vein ending at the 

dental sinus. All Rosaceae that have teeth and (or) have 

deeply dissected laminae tend to have moderately strong 

to marked sinal braces, many of which originate from the 

apical sides of secondary or external secondary veins; this 

syndrome may be homologous to the apical branch in a 

typical cunonioid tooth. 

The lamina in many rosaceous taxa is deeply 

dissected and may be totally pinnatisect but apparently 

rarely, if ever, truly compound (Wolfe and Wehr, 1988). 

This applies to rosaceous groups that are fundamentally 

pinnately veined (Spiraeoideae and Pyroideae ), as well as 

to some actinodromous groups (Rosoideae, in part). 

Whether Rosaceae are primitively pinnately veined or 

actinodromous is, as yet, unresolved, but the most 

primitive extant members of Spiraeoideae, as well as of 

Pyroideae and Prunoideae, are simple-leaved. In at least 

one instance, evolution of pinnatisect leaves from a 

pinnatifid ancestor has been inferred (Wolfe and Wehr, 

1988). 

Somewhat transverse, A-0 tertiary veins are 

typical of many Rosaceae. This architecture could link 

Rosaceae with the iteid rosids (see discussion of Jamesia), but the numerous composite intersecondary veins 

and typical brochidodromy in Rosaceae are not iteid 

characters. Finally, most Rosaceae are markedly 

stipulate, and the stipules tend to be persistent. 

Subfamily SPIRAEOIDEAE 

Genus ELEIOSINA Raflnesque 

Elelosina praecondnna (Cockerell) Sc:bom & Wolfe, n. comb. 

Plate 7, figures 1-5 

Solidago praeconcinna Cockerell, 1933, p. 72, fig. 1. 

Salix creedensis Axelrod, 1987, p. 105, pl. 18, fig11. 5, 6; pl. 20, fig11. 2, 4-6. 

Cercocarpus holmesii auct. non (Lesquereux) Axelrod, 1944. Axelrod, 1987 

(part), p. 124. 

Peraphyllum septentrionalis auct. non Daphne septentrionalis (Lesquerex) 

MacGinitie. Axelrod, 1987, p. 132, pl. 24, fig11. 12, 13. 

Wolfe and Schorn (1989) designation: Eleiosina sp. 

Description.-Leaves alternate, simple; apparently 

stipulate; shape symmetric, elliptic to narrowly elliptic; 

base symmetric and acute, apex acute to acuminate and 

has a short (rarely elongated) mucro; length 2.1->7.2 

em, width 0.5-1.1 em; venation pinnate; midrib weak; 

7-10 pairs of secondary veins, markedly decurrent 

(especially basally) then extending at angles of 200-300 to 

midrib, thin, straight to typically slightly convex, offset 

where a major branch originates, weakly brochidodromous to reticulodromous; intersecondary veins simple, 

common, some joining subjacent secondary vein; tertiary 

veins thin, R-R, widely spaced, reticulate; fourth-order 

veins forming imperfect, thin, irregularly polygonal areales; veinlets thin, dendritically branching; external 

secondary veins three to four per secondary vein, 

originating at acute angles, some originating close to 

midrib, forming loops that are elongated parallel with 

margin, some reticulodromous; margin entire, thin; 

petiole absent. 

Systematic Descriptions and Discussions 23 
Discussion.-The critical characters of the Creede 

leaves here grouped under Eleiosina praeconcinna are 

(1) an entire margin that is unthickened and has no 

fimbria! vein, (2) a glandular, slightly mucronate apex, 

(3) secondary veins that are markedly decurrent and 

extend at very low angles relative to the midrib, (4) 

secondary veins that are thin and are offset at points of 

origination of major branches, (5) secondary veins that 

are weakly brochidodromous or that may not loop (that 

is, some secondary veins are reticulodromous), (6) 

numerous external secondary veins that originate at low 

angles, (7) some external secondary veins that originate 

close to the midrib, (8) external secondary veins that are 

weakly brochidrodromous or reticulodromous, (9) intersecondary veins that extend over half the distance to the 

margin and that can merge with a secondary vein so as to 

isolate a large part of an intercostal area, and (10) widely 

spaced tertiary veins that form an irregularly branched 

reticulum. The holotype of E. praeconcinna, moreover, 

has alternate leaves, and stipules appear to be present. 

Of the previously listed characters, Salix leaves 

typically lack characters 3, 5, and 7-10. Even in gross 

characters, the supposed "match" for Salix creedensis, S. 

geyeriana Anderss., markedly differs and has leaves that 

typically have a few to several salicoid teeth, markedly 

brochidodromous to eucamptodromous secondary veins, 

and an acute, non-mucronate apex. 

Peraphyllum leaves are perhaps closer than Salix 

leaves to Eleiosina praeconcinna, but Peraphyllum leaves 

(pl. 7, fig. 7) typically do not have decurrent secondary 

veins or a mucronate apex, the secondary veins and 

external secondary veins are markedly brochidodromous, 

the secondary veins are not offset at point of origination 

of major branches, and external secondary veins originate 

close to the margin. 

An entire margin, marked decurrency of secondary 

veins, and poorly organized venation are generally 

primitive characters in the dicotyledons and are 

particularly characteristic of some taxa of Magnoliidae 

(Hickey and Wolfe, 1975; Hickey and Doyle, 1977). 

Search has revealed no magnoliid that has a mucronate, 

apparently glandular apex associated with these other 

characters, and the fossils show no indication of oil cells 

characteristic of Magnoliidae. In subclasses other than 

Magnoliidae, these primitive characters are rarely found 

in combination. In Crossosoma (Crossosomataceae), a 

putative primitive member of Dilleniidae (but placed by 

some near Rosaceae), these primitive characters are 

found but in combination with marked festooned brochidodromy. In Eleiosina leaves (pl. 7, fig. 6), however, are 

the 10 characters, including the 3 primitive characters, 

listed previously for the fossils, although the Creede 

24 Spermatopsida of Oligocene Creede Flora, Colorado 

leaves typically do not have as pronounced a mucro (but 

see pl. 7, fig. 3) as the leaves of the four extant species of 

Eleiosina examined and have an elliptic rather than an 

obovate shape. 

The fossils from floras other than Creede that 

Axelrod (1987) assigned to Peraphyllum septentrionalis 

are here rejected as belonging to Eleiosina praeconcinna. 

These other fossils do not have a mucronate apex or 

decurrent secondary veins, have markedly brochidodromous secondary and external secondary veins, and have 

secondary veins that maintain a low origination angle in 

the apical region (pl. 7, fig. 8). Proper taxonomic 

disposition of these other fossils awaits more detailed 

examination and analysis. 

Nomenclature.-Axelrod (1987) synonymized Solidago praeconcinna under Cercocarpus holmesii. Total 

absence of teeth, thin secondary veins that are widely 

spaced, and presence of a thin midrib on Cockerell's 

material readily differentiate S. praeconcinna from C. 

holmesii (see later under C. henricksonii). 

Holotype.-UCMG 18536. 

Hypotypes.-VCMP 7496-7506,7386,7388, 7389; 

UCMG 27387A, B. 

Genus ELEOPOLDIA Schorn & Wolfe, n. gen. 

Diagnosis.-Leaves simple; apparently estipulate; 

not pubescent; pinnatifid, typically three (some two or 

four) lobes on either side of lamina, with basal part of 

lobe decurrent on midrib and including a weakly decurrent vein that enters basal part of lobe; most basal lobes 

also decurrent to base of petiole; secondary laminar 

segments pinnatifid, as many as three lobes; some 

tertiary laminar segments have a tooth on basal side; 

margin otherwise entire and not revolute; venation pinnate; fourth-order veins forming distinct, irregularly 

polygonal areoles; branching freely ending veinlets. 

Type species.-F allugia lipmanii Axelrod. 

Nomenclature.-This genus is named for Estella 

Leopold, in recognition of her many contributions to the 

Tertiary palynology, paleobotany, and historical plant 

geography of the Rocky Mountains. 

Eleopoldia lipmanii (Axelrod) Schorn & Wolfe, n. comb. 

Plate 8, figures 1, 3, 5, 8, 9; pL'lte 13, flgure 3 

Fallugia lipmanii Axelrod, 1987 (part, typic), p. 129, pl. 29, figs. 1-9, 11, 12. 

Junipems gracillensis Axelrod, 1987, p. 100, pl. 16, figs. 10-12. 

Wolfe and Schorn (1989) designation: aff. Luetkea sp. 

Description.-Leaves simple; apparently estipulate; 

glabrous; shape symmetric, orbiculate to oblate, pinnatifid; base symmetric, decurrent, apex acute to narrowly 

obtuse; length 1.0-4.0 em, width 1.1->4.0 em; venation 

pinnate; midrib stout; two to five pairs of secondary veins, 

originating decurrently then extending &t angles of 
30°-80°, straight to broadly convex or broadly concave, 

entering lobe along apical side of lobe, craspedodromous; two to three external secondary veins originating 

at low angles from basal side of secondary veins, one to 

two from apical side, craspedodromous; sinuses of 

tertiary laminar segments braced by a minor branch from 

subjacent external secondary veins, branching at sinus, 

these branches extending parallel with sides of sinus; veins 

about same thickness as external secondary branches 

originating decurrently from midrib, extending into lobe 

subjacent to secondary vein; fourth-order venation 

forming somewhat imperfect, irregularly polygonal areales; veinlets dendritically branching; one to four pairs of 

secondary laminar segments, decurrent along midrib 

narrowly elliptic to obovate, narrowly obtuse to acut~ apices, pinnatifid; two to three narrow tertiary laminar 

segments on basal side of secondary laminar segments, 

one to two on apical side; some tertiary laminar segments 

wit~ one tooth; sinuses acute to narrowly rounded; 

petiole 0.2-2.1 em long. 

Discussion.-Leaves that are pinnatifid and have 

strongly decurrent secondary laminar segments occur in 

several extant genera of Rosaceae, includingFa/lugia and 

Purshia. Leaves of Fal/ugia (pl. 8, fig. 7), however, are 

stipulate, strongly pubescent, and strongly revolute and 

have secondary laminar segments that are rarely lobed (if 

lobed, teeth are absent on the lobes). Leaves of Purshia 

ar~ shed without stipules (and thus could appear to be 

estipulate ), and some laminae can have a rare lobe on a 

secondary laminar segment, but the strong pubescence 

and strongly revolute margin also eliminate this genus 

from consideration. 

Three extant genera of Rosaceae, all herbaceous, 

appear as the closest extant relatives of Eleopoldia: 

Luetkea, Geum, and Sanguisorba. Luetkea has estipulate, 

n?npu?~scent, nonrevolute leaves (pl. 8, fig. 2) that are 

pmnatifid and that have pinnatisect secondary laminar 

segments; typically, however, only a single pair of lateral 

lobes is present, the tertiary laminar segments never have 

te~th, and the secondary laminar segments have a single 

pau of lobes. Geum has leaves (pl. 8, fig. 6) that have as 

many as two lobes on the secondary laminar segments 

and have areolation as in Eleopoldia, but these extant 

leaves are stipulate, pubescent, and pinnatisect. Sanguisorba has leaves (pl. 8, fig. 4) that have several lobes 

on the secondary laminar segments and lack pubescence, 

but the leaves are stipulate and pinnatisect and the 

secondary laminar segments lack teeth. We emphasize, 

however, that absence of stipules on mature leaves that 

enter the fossil record could also result from the stipules 

being deciduous. 

Eleopoldia was probably a woody plant (Wolfe and 

Schorn, 1989); leaves of terrestrial dicotyledonous herbs 

rarely enter the fossil record (they generally do not 

dehisce from the plant) and then never abundantly. 

Because Eleopoldia has the greatest similarity to leaves 

of Luetkea, we suggest that Eleopoldia may have given 

rise to this extant genus; the differences between leaves 

of the two genera appear primarily as reductions: fewer 

secondary laminar segments, fewer tertiary laminar 

segments, and absence of teeth in Luetkea. Luetkea also 

typically has more specialized venation in that the points 

of origination of the secondary veins and the major 

branches of these veins are close to sinuses and the 

secondary veins tend not to be decurrent. We would thus 

consider Eleopoldia to be a member of Spiraeoideae, as 

is Luetkea. The relationship of Eleopoldia to Geum and 

Sanguisorba is probably more distant; the absence of the 

plesiomorphic (for Rosaceae) stipulate and pubescent 

conditions suggests that the Eleopoldia-Luetkea line 

diverged from ancestral stock that could have given rise 

to these other two extant genera (and possibly other 

members of Rosoideae ). 

Nomenclature.-The epithet lipmanii, which is 

based on a large suite of well-preserved specimens, 

should be selected over gracil/ensis, which is based on a 

few, poorly preserved specimens. 

Holotype.-VCMP 7767. 

Paratypes.-VCMP 7761-7765, 7768-7780, 

7782-7785. 

Hypotypes.-UCMP 7215, 7445, 7446; UCMG 

19786,34068,34069. 

Genus STOCKEYA Wolfe & Wehr 

Stockeya creedensis (R. W. Brown) Wolfe & Wehr 

Stockeya creedensis (R.W. Brown) Wolfe & Wehr, 1988, p. 193, figs. 14, 15, 

32-36, 50-52. 

Chamaebatiaria creedensis R. W. Brown, 1937, p. 177, pl. 57, figs. 8, 9. Axelrod, 

1987, p. 127, pl. 27, figs. 1-14. 

Wolfe and Schorn (1989) designation: Stockeya creedensis. 

Discussion. -Stockeya creedensis was extensively 

discussed by Wolfe and Wehr (1988). Stockeya, of which 

S. creedensis is the genotype, is considered probably to be 

ancestral to the extant Chamaebatiaria. 

Lectotype.-USNM 39651. 

Hypotypes.-UCMP 7715-7742; UCMG 18646, 

34057, 34058. 

Genus HOWDISCUS Maximowicz 

Holodiscus stevenii Schorn & Wolfe, n. sp. 

Plate 9, figures 3, 4, 7, 8 

Holodiscus hameyensis auct. non (Arnold) Axelrod, 1950. Axelrod, 1987 (part), 

p. 130, pl. 30, figs. 1-4. 

Holodiscus idahoensis auct. non Chaney & Axelrod nom. nud. Axelrod, 1987, p. 

131, pl. 30, figs. 5-11. 

Phyllites sp. Knowlton, 1923, p. 191, pl. 43, figs. 13, 15. 

Rosa hilliae auct. non Lesquereux. Axelrod, 1987, p. 137, pl. 31, fig. 11. 

Wolfe and Schorn (1989) designation: Holodiscus sp. 

Systematic Descriptions and Discussions 25 
Description. -Leaves simple; apparently estipulate; 

shape symmetric, elliptic to (in larger specimens only) 

ovate; base symmetric, decurrent, apex acute to narrowly 

rounded; length 1.1->4.0 em, width 0.5-1.7 em; venation pinnate; midrib moderate; three to four pairs of 

secondary veins, originating typically at angles of 20°-30° 

(rarely greater), straight to slightly convex (rarely 

concave), extending through medial part of tooth, craspedodromous and flanked on both sides by weak veins that 

converge on apex of tooth; rarely one to two strong veins 

originating at higher angles from midrib and basal to 

most basal secondary veins, brochidrodromous; tertiary 

veins A-A to R-A, widely spaced, reticulate; fourthorder veins forming small, imperfect, irregularly 

polygonal areoles; veinlets dendritically branching; 

external secondary veins originating acutely, few, 

thinning, brochidrodromous or rarely craspedodromous 

(if subsidiary tooth present); dental sinuses braced by 

external veins that originate from both suprajacent and 

subjacent secondary veins and merge at sinus; margin has 

as many major teeth as secondary veins, A-1 or D-1 

(rarely A-4), typically no subsidiary teeth; subsidiary 

tooth, if present, on second most basal major tooth; 

sinuses acute to narrowly rounded. 

Discussion.-Both Holodiscus and Cercocarpus can 

have teeth that are weakly rosoid and confined to the 

apical half of the lamina. In Holodiscus leaves, the most 

basal secondary veins are placed away from the margin, 

brochidodromous external secondary veins are present, 

and the dental sinuses have conspicuous braces; stipules, 

if present, are deciduous, and thus fossil foliage will 

appear estipulate. In Cercocarpus leaves, the most basal 

secondary veins are placed close to the margin, typically 

no external secondary veins are present, and the dental 

sinuse~ have very reduced braces; stipules are present. 

On these criteria, the leaves referred by Axelrod (1987) 

to Holodiscus represent, in large part, Holodiscus. 

Extant Holodiscus can be separated into two 

groups based on decurrency of the lamina and number of 

secondary teeth. In one group, a distinct, naked petiole 

and numerous secondary teeth are present: H. discolor 

(Pursh) Maxim., to which the larger Creede leaves were 

compared, H. dumosus Nutt., and H. glabrescens 

Greenm. In the second group, the lamina is strongly 

decurrent along the petiole and secondary teeth are few 

or absent: H. boursieri (Carr.) Rehd. and H. microphyllus 

Rydbg. The smaller Creede leaves were likened to H. 

dumosus, H. glabrescens, and H. microphyllus. Both large 

and small leaves of H. stevenii fall into the second group, 

as do Neogene leaves of H. fryi Wolfe. In contrast, the 

type of H. harneyensis (Arnold's, 1937, pl. 4, fig. 3) falls 

into the first group. The leaf referred to H. hameyensis by 

Axelrod (1966, pl. 16, fig. 10; erroneously synonymized 

with Crataegus cuprovallis by Wolfe, 1977) has the laminar decurrency of the second group, but presence of 

26 Spermatopslda of Oligocene Creede Flora, Colorado 

secondary teeth indicates a relationship to the first group; 

the Copper Basin taxon may represent an ancestor to 

both extant groups of Holodiscus. Smith's (1941, pl. 12, 

fig. 1) specimen of Cercocarpus cuneatus Dorf (what was 

to become the type of H. idahoensis) has an incomplete 

base, but the few secondary teeth suggest membership in 

the second group. 

The sole basis for separating the Creede specimens 

assigned to Holodiscus harneyensis from those assigned 

to H. idahoensis was size (minimal length 4 em versus 

maximal length 2 em, respectively (Axelrod, 1987, p. 

130-131)), although cited but unfigured specimens ofthe 

larger leaved H. harneyensis included two only 2.5 em 

long. Note that the three localities that produced the five 

larger leaves of H. stevenii are localities that produced 

samples inferred by Wolfe and Schorn (1989) to 

represent mesic, closed-canopy forest (Axelrod's 

localities 571-A, 571-B, and 574-A), whereas the 103 

smaller and more typical leaves of H. stevenii occur in 

samples that were inferred to represent the full range of 

Creede vegetation, from mesic forest to chaparral. 

The specimen previously assigned to Rosa hilliae 

(pl. 9, fig. 8) appears at first glance to be different from 

leaves of H. stevenii; however, in details of laminar 

venation and dental morphology, this specimen is like H. 

stevenii. What gives the specimen assigned to R. hilliae a 

different appearance are the convex sides in the basal 

part of the lamina, strong veins that originate from the 

midrib basal to the most basal pair of secondary veins, 

and the asymmetric base. Some specimens of the Creede 

Holodiscus, however, have convex sides basally (Axelrod, 

1987, pl. 30, figs. 3, 6), and some have strong veins basally 

(Axelrod, 1987, pl. 30, fig. 6). The base of the specimen 

of R. hilliae, moreover, is folded to give a falsely 

asymmetric appearance. 

If, as considered by Axelrod (1987), the Creede 

Holodiscus and H. fryi were conspecific, then H. fryi 

would be the oldest validly published name for the 

Creede material. In H.fryi, however, (1) the lamina has a 

length to width ratio of about one to one in contrast to 

greater than two to one ratio in H. stevenii, (2) the most 

basal lobes flare abmedially in contrast to their apical 

orientation in H. stevenii, (3) the secondary veins 

originate at angles of 40°-50° in contrast to less than. 

30° in H. stevenii, and (4) if subsidiary teeth occur, they 

are on the most basal lobes in contrast to being on the 

second most basal lobes in H. stevenii. The characters 

that separate leaves of H. ftyi from leaves of H. stevenii 

are primarily the same characters that separate leaves of 

H. boursieri from leaves of H. microphyllus. 

Nomenclature.-As just discussed, the holotype of 

Holodiscus idahoensis is incomplete and lacks a character 

critical in taxonomic partitioning of Holodiscus foliage. 

More importantly, Smith (1941) presented no 

description of her specimen of Cercocarpus cuneatus, 
and Chaney and Axelrod (1959) failed to present either 

description or diagnosis; Axelrod's (1987) cursory 

description, whether judged adequate, was stated to be 

based on the Creede leaves only and cannot be construed 

as validating H. idahoensis, which must be considered 

under the ICBN an illegitimate name. As also just 

discussed, H. fryi represents a taxon distinct from the 

Creede Holodiscus, which thus represents a new species. 

We take pleasure in naming this distinctive Holodiscus 

for Thomas Steven, who has made numerous 

contributions to the geology of the Creede area. 

Holotype.-UCMG 34070. 

Paratypes.-VCMP 7705, 7786-7789, 7791-7816, 

7821; UCMG 34071, 34072. 

Subfamily PYROIDEAE 

Genus SO RBUS Linnaeus 

Sorbus potentilloldes (Knowlton) Axelrod 

Plate 9, figure 5 

Sorbus potentilloides (Knowlton) Axelrod, 1987, p. 138, pl. 31, figs. 2-6, 8. 

Phyllites potentilloides Knowlton. 1923, p. 191, pl. 43, fig. 14. 

Wolfe and Schorn (1989) designation: Sorbus potentilloides. 

Description.-Leaves odd-pinnate; at least nine 

leaflets; leaflet shape symmetric, lorate; base rounded to 

acute and some strongly asymmetric, apex acute to 

narrowly obtuse; length 1.3-3.5 em, width 0.4-0.8 em; 

venation pinnate; midrib moderate; seven to nine pairs of 

secondary veins, originating at angles of 30°-50°, straight 

to slightly convex, rarely branching before entering teeth, 

entering teeth medially, flanked by weak converging 

veins, craspedodromous; tertiary veins typically R-R, 

branching, reticulate; fourth-order veins forming welldeveloped, almost quadrangular areoles; veinlets 

typically once branching; dental sinuses braced by 

tertiary veins that originate from either subjacent or 

suprajacent secondary vein; teeth typically one per 

secondary vein but some leaflets have no teeth in basal 

third oflamina; teeth typically B-1, basal side about three 

times as long as apical side; sinuses angular; apetiolulate 

or with petiolule < 0.1 em long. 

Discussion-Although Becker's (1962) Sorbus 

charcharodonta was synonymized with S. potentilloides by 

Axelrod, the leaflets of S. charcharodonta have rounded 

A-1 teeth and more numerous secondary veins that 

originate at high ( > 60°) angles. The relationships of the 

various described species of Sorbus in the North 

American Tertiary are poorly known, however, and the 

fossil material of this genus is in need of thorough 

revision. The Creede laminae, which have no subsidiary 

teeth, appear related to the extant S. aucuparia L. (Axelrod, 1987, p. 139). 

Holotype .-USNM 36539. 

Hypotypes.- UCMP 7822-7828; UCMG 19773. 

Genus CRATAEGUS Toumefort ex Linnaeus 

Crataegus creedensls Axelrod 

Plate 9, figure 6 

Crataegus creedensis Axelrod, 1987, p. 128, pl. 28, figs. 1-3. 

Wolfe and Schorn (1989) designation: Crataegus creedensis. 

Description.-Leaves simple; apparently estipulate; 

shape symmetric, ovate to elliptic, pinnatifid to almost 

pinnatisect; base symmetric, cuneate, apex acute to 

narrowly rounded; length 1.4- > 2.9 em, width 1.0-2.2 

em; venation pinnate; midrib moderate; three to five 

pairs of secondary laminar segments, ovate to typically 

elliptic to obovate, apex acute to narrowly obtuse, base 

strongly decurrent to subjacent secondary laminar 

segment; secondary veins originating at angles of30°-50°, 

entering laminar segment either medially or toward 

apical margin of segment, straight to slightly concave, 

craspedodromous; one to two veins thinner than 

secondary veins originating basal to secondary veins and 

extending parallel with secondary vein; tertiary veins in 

medial (apical) laminar segment R-R to A-R, reticulate; 

tertiary veins in lateral secondary segments R to A and 

forming brochidodromous loops or rarely craspedodromous; fourth-order veins forming imperfect, 

quadrangular to pentagonal areoles; veinlets typically 

once branched; lobal sinal bracing formed by a weak 

intersecondary vein that branches at sinus, each branch 

weak and extending a short distance close to sinal 

margin; each secondary segment typically has one tooth 

on basal side near apex of segment; sinuses angular; 

petiole 0.2-0.6 em long. 

Discussion-Most leaves of Crataegus creedensis 

are almost pinnatisect, a rare condition that occurs in 

some leaves of some species of extant Crataegus. No 

extant species of the genus appears closely related to C. 

creedensis. Although compared by Axelrod (1987) to 

extant species such as C. azarolus L., C. creedensis has 

weak lobal sinal braces (in C. azarolus the bracing vein 

and its branches are thick and extend half or more the 

distance to the apex of the secondary laminar segments) 

and thin but straight secondary veins (in C. azarolus, the 

thick secondary veins are offset at the points of 

origination of major branches). 

Holotype.-UCMP 7259. 

Paratypes.-UCMP 7754-7758, 7760. 

Subfamily ROSOIDEAE 

Genus POTENTILLA Linnaeus 

Potentilla creedensis (Axelrod) Schorn & Wolfe, n. comb. 

PL'lte 10, figure 7; pL'lte 11, figure 1 

Ranunculus creedensis Axelrod, 1987, p. 109, pl. 21, fig. 6. 

Wolfe and Schorn (1989) designation: Potentilla sp. 

Description. -Leaves simple; stipulate; tripalmatisect and palmatifid; the three pairs of lateral primary 

Systematic Descriptions and Discussions 27 
laminar segments doubly palmatifid; shape symmetric, 

widely ovate; base deeply cordate, apex acute; length 

1.5-3.5 em, width 2.0-3.8 em; lateral primary laminar 

segments ovate, palmatifid, base cuneate; venation 

perfectly actinodromous; midrib moderate; the six lateral 

primary veins moderate, slightly concave; three to four 

pairs of secondary veins originating from midrib and 

from more apical pair of lateral primary veins and one to 

two pairs from more basal pairs of lateral primary veins 

at angles of about 30°, typically concave, entering teeth 

medially, flanked by weak veins, craspedodromous; 

dental sinal braces formed by external vein that 

originates from subjacent secondary vein; higher order 

venation obscure; secondary laminar segment have one 

to two A-1 teeth on basal side and zero to one on apical 

side; sinuses angular; petiole 0.9-1.7 em long. 

Discussion.-Ranunculaceae have chloranthoid 

teeth (Hickey and Wolfe, 1975), whereas the Creede 

specimen assigned to Ranunculus has weakly developed 

rosoid teeth. Palmatisect to palmatifid foliage in 

association with weakly developed rosoid teeth is 

particularly common in Potentilla. 

Potentilla creedensis is actinodromous and is thus 

generally related to the many extant species that are 

palmatisect. Most of these species, however, have laminae that are pentapalmatisect, in contrast to P. creedensis 

laminae, all of which are tripalmatisect and have lateral 

segments twice palmatifid; this condition has not been 

observed in extant Potentilla. In some Arctic tundra 

species (for example, P. noroegica L. and P. groenlandica 

Malte ), laminae are tripalmatisect; the most basal lobes 

of the lateral segments are entered by veins that originate 

basally and could thus be regarded as reduced lateral 

primary veins, but the lobes are about the same size as 

more apical secondary laminar segments. In P. subquinata leaves, the laminae are also typically tripinnatisect as 

in the Arctic taxa, but some are pentapalmatisect; in the 

pentapalmatisect laminae, the most basal major veins of 

the most apical pair of lateral segments are secondary 

veins, suggesting that indeed the most basal major veins 

of tripalmatisect leaves are reduced primary veins. 

In this context, foliage of the Creede Potentilla 

appears to represent a stable intermediate condition 

between the exclusively tripalmatisect Arctic species and 

the exclusively pentapalmatisect species. The Arctic 

leaves, however, have untoothed, robust secondary laminar segments, in contrast to the toothed, filiform 

secondary laminar segments of Potentilla creedensis. The 

apparent relationship of the Creede Potentilla to the 

Arctic tundra taxa may indicate that some Arctic tundra 

lineages originated in middle-latitude, high-altitude 

regions, a concept held by some plant geographers. 

Holotype.-VCMG 19757. 

Hypotypes.- USNM 422590; HUBM 62425 

28 Spermatopsida of Oligocene Creede Flora, Colorado 

Genus CERCOCARPUS Humboldt, Bonpland, & Kunth 

Cercocarpus henricksonii Schorn & Wolfe, n. sp. 

Plate 10, figures 3--6, 15, 16 

Cercocarpus holmesii auct. non (Lesquereux) Axelrod, 1944. Axelrod, 1987 

(part), p. 124, pl. 26, figs. 1-14. 

Planera myricaefolia auct. non (Lesquereux) Cockerell. Knowlton, 1923, p. 188, 

pl. 43, figs. 16, 17. 

Wolfe and Schorn (1989) designation: Cercocarpus sp. 1. 

Description.-Leaves simple; stipulate; shape 

symmetric, very narrowly elliptic or oblanceolate to 

narrowly oblanceolate; base symmetric, acute to cuneate, 

apex acute to narrowly obtuse; length 2.7-6.0 em, width 

0.3-1.4 em; venation pinnate; midrib stout; 6-12 pairs of 

secondary veins, originating at angles of 10°-20°, typically 

straight, the basal one to three pairs brochidodromous, 

the more apical pairs entering teeth medially, flanked by 

two weak converging veins, craspedodromous; tertiary 

veins R-R, closely spaced; fourth-order veins forming 

with tertiary veins small, well-developed, quadrangular 

areoles; veinlets unbranched; teeth simple, typically 

confined to apical half of lamina, A-1 or B-2, apical side 

very short relative to basal side and some teeth are very 

appressed; sinuses angular; margin revolute; petiole 

0.2-0.6 em long. 

Discussion.-The architecture of these leaves is 

consistent with assignment to Cercocarpus (see previous 

discusssion ofHolodiscus). Particularly note the stipulate 

condition (pl. 10, fig. 3). 

Cercocarpus bea-anniae Beck., which was 

synonymized with the Creede material by Axelrod 

(1987), is based on leaves that have only three to five 

pairs of secondary veins, a convex curvature to the 

secondary veins, and a few teeth confined to the most 

apical part of the lamina. Although conceivably Becker's 

(1962) leaves could represent extreme variants of the 

Creede species, none of the Creede leaves of C. henricksonii that we have examined combine the characters on 

which C. bea-anniae was founded. 

The Florissant Cercocarpus myricaefolius (Lesq.) 

MacG. may be related to C. henricksonii. The Florissant 

leaves, however, typically have large, sharp D-2 or D-3 

teeth that occur in the basal half of the lamina. 

The presumed achenes assigned to Cerococarpus 

holmesii by Axelrod (1987) present a problem rather 

than giving confirmatory evidence as to the presence of 

the genus in the Creede flora. Some specimens have 

evidence of hairs on the elongated styles, which would 

indicate Cercocarpus. However, the specimen illustrated 

by Axelrod (1987, pl. 29, fig. 10) as Fallugia lipmanii may 

represent the same taxon as the isolated achenes 

assigned to Cercocarpus. The specimen (pl. 13, figs. 1, 2) 

has three curved, long styles (preservation is too poor to 

determine pubescence) that emanate from an apparently 

striated structure that may represent the hypanthium 

tube. Two presumably indehiscent structures, also with 
curved, long styles, cross the hypanthium tube, and the 

presumed achenes are almost exactly the same length as 

the hypanthium tube. These other two structures appear 

as if they were partially removed from the hypanthium 

tube at the time of deposition; in all preserved characters, 

these two structures are identical to the achenes assigned 

to Cercocarpus from other Creede localities; the slightly 

smaller size of the achenes and shorter styles than in the 

other isolated achenes is probably related to being 

immature. This specimen was collected at Axelrod's 

locality 573-A, where 131 leaves of our C. henricksonii 

but only 42 leaves of other angiosperm taxa (including 25 

of Populus, 7 of Ribes, and 3 of Eleopoldia) were censused, and this association suggests that this specimen 

would more probably represent the same taxon as the 

Cercocarpus leaves rather than the Eleopoldia ( = Axelrod's Fallugia) leaves. On the other hand, this pentacarpellary structure cannot represent true Cercocarpus, 

which is exclusively unicarpeiiate, although it is 

associated with Cercocwpus-like leaves and isolated 

achenes have been accepted as Cercocarpus. 

This raises the possibility that, if the leaves and 

reproductive structures represent the same taxon, Cercocarpus henricksonii (and possibly the Florissant C. myricaefolius) represents a genus ancestral to Cercocarpus, 

and that five carpels (which are surely primitive in 

Rosaceae) have been reduced to one. On the other hand 

the presence in the Creede of C. nanophyllus, which 

represents the distinct, specialized C. ledifolius clade, 

would indicate Cercocarpus had already originated prior 

to Creede time. "Cercocarpus" henricksonii would, in this 

context, not be a species ancestral to any extant true 

Cercocarpus but only a lingering species of the ancestral 

genus. Additional, better preserved specimens and 

additional evidence from more associations with the 

Cercocarpus-like foliage would test this hypothesis. The 

possibility of the validity of the hypothesis suggests that 

caution should be exercised before attempting to relate 

C. henricksonii to any extant species of Cercocarpus. 

Although the leaves of Cercocarpus henricksonii do 

not appear to differ in any major architectural features 

from extant Cercocarpus. the stipules are free (pl. 10, fig. 

3), whereas in extant Cercocarpus (as well as in C. 

nanophyllus; pl. 10, fig. 14), the stipules are adnate to the 

petiole except at their apices. Additional, detailed 

architectural comparisons between extant and fossil 

foliage are needed. 

Nomenclature.-The types of Lesquereux' (1887) 

Crataegus holmesii, which form the basis for Cercocarpus 

holmesii (Lesq.) Axelr., are here refigured (pl. 10, figs. 1, 

2, 8, 9). These specimens were coiiected from a locality at 

Silver Cliff, Colo., and assignment of the Creede leaves of 

toothed Cercocarpus to C. holmesii is dubious at best. 

One of the types of C. holmesii (pl. 10, fig. 1) has no 

indication of secondary veins and is entire-margined; this 

specimen cannot be generically or familially determined. 

The margin of the second specimen can be interpreted as 

either an original margin or one created by tearing along 

veins; the specimen cannot be determined. The third 

specimen is a fragment of a lamina that had at least three 

pairs of secondary veins that originated at very low 

angles; no teeth and no higher order venation are 

preserved. The fourth specimen is a fragment of the 

apical portion of a lamina; the secondary veins originate 

at very low angles and appear to terminate in very 

appressed teeth, but greater detail is not preserved. The 

fourth specimen could well represent the same species as 

C. henricksonii but could equally well represent the same 

species as C. bea-anniae or even some as yet undescribed 

species. The types of C. holmesii are too fragmentary and 

too poorly preserved to determine their affinities with 

any degree of certainty. 

Axelrod (1987) erred in designating as lectotypes 

for Cercocarpus holmesii two specimens (for a given 

taxon, only one lectotype can be designated) and also 

erred in designating USNM 36541 and 36542. These two 

Creede specimens are those illustrated by Knowlton 

(1923) as Planera myricaefolia; under the ICBN, a lectotype for Lesquereux' (1887) C. holmesii must come from 

the original type suite. We herewith designate as lectotype of Crataegus holmesii Lesq. USNM 313299 (pl. 10, 

fig. 9). We take pleasure in naming this species for James 

Henrickson, who has contributed greatly to 

understanding the phylogeny and systematics of extant 

Rosaceae. 

Holotype.-VCMG 34073 (counterpart USNM 

422592). 

Paratypes.-VSNM 36541, 36542; UCMP 

7679-7713; UCMG 34074-34077. 

Cercocarpus nanopbyllus Schorn & Wolfe, n. sp. 

Plate 10, figures 12-14 

Cercocarpus linearifolius auct. non (Lesquereux) Axelrod, 1950. Axelrod, 1987, p. 

126, pl. 28, figs. 5-11. 

Wolfe and Schorn (1989) designation: Cercocarpus sp. 2. 

Description.-Leaves simple; stipulate; shape 

symmetric, narrowly to very narrowly elliptic, typically 

falcate; base symmetric, acute to obtuse, apex narrowly 

obtuse to acute; length 1.0--2.5 em, width 0.2--0.3 em; 

venation pinnate; midrib massive (as much as 15 percent 

of the width of lamina); 15-30 pairs of closely spaced 

secondary veins, originating at angles of 

50°-(typically) > 70°, straight, forming markedly brochidodromous loops; abmedial segments of loops tending to 

form an intramarginal vein; numerous composite intersecondary veins, some of which extend to abmedial 

segment of loop and thus may not be clearly 

distinguished from secondary veins; tertiary veins 

typically A-R, forming well-developed, typically 

Systematic Descriptions and Discussions 29 
quadrangular areoles; veinlets few, unbranched or once 

branched; margin entire, strongly revolute; petiole < 0.1 

em long. 

Discussion. -Entire-margined laminae that are 

reduced in size, such as those just described, typically 

have reduced venation, which makes their identification 

difficult. Characters may then have to be employed that 

are of minor taxonomic significance. For example, the 

markedly revolute margin of these leaves is not of major 

taxonomic significance and can be found in Ledum in 

Ericaceae, as well as in Cercocarpus. Ledum has, 

however, as do most Ericaceae, reticulodromous venation and, like all Ericaceae, is estipulate. The Creede 

leaves cannot, therefore, represent Ericaceae. 

These Creede leaves also have closely spaced A-R 

tertiary veins and numerous composite intersecondary 

veins. This architecture, although not confined to 

Rosaceae, is particularly common in the family. 

Combined with the stipulate condition, the assignment by 

Axelrod (1987) of these leaves to Rosaceae based on 

resemblance to the extant Cercocarpus ledifolius Nutt. 

appears warranted, although we emphasize the 

somewhat tenuous nature of the generic and familial 

assignment. 

Although Cercocarpus nanophyllus is apparently 

related to the extant C. ledifolius, leaves of the two 

species are readily distinguished. In C. ledifolius leaves 

the secondary veins typically originate at an angle of less 

than 45°, and the abmedial segments of the secondary 

vein loops are aligned to form an intramarginal vein only 

in the apical region of the lamina. 

Nomenclature.-Axelrod (1987) included the 

Creede material here described as Cercocarpus nanophyllus in C. linearifolius, which is based on Lesquereux' 

(1887) Andromeda linearifolia from Silver Cliff, Colo. 

Because C. linearifolius has been cited as occurring 

widely in the Neogene of western North America (Axelrod, 1950, 1956, 1987), as well as in the Creede flora, we 

reexamined the type material of this taxon. 

The extant type suite of Andromeda linearifolia 

includes only two specimens, one represented by part and 

counterpart. One specimen is a complete lamina but has 

only vague indications of secondary veins (pl. 10, fig. 11 ); 

assignment of this specimen to the angiosperms could be 

questioned. The second specimen (pl. 10, fig. 10), which 

lacks base and apex, has a few, widely spaced secondary 

veins that originate at very low angles (about 20°), an 

exceedingly thick midrib (the width of the midrib is about 

20 percent of the width of the lamina), and a markedly 

revolute margin (as does the first specimen). When 

contrasted to the closely spaced secondary veins that 

originate at high angles in Cercocarpus nanophyllus, A. 

linearifolia is clearly distinct. We herewith designate as 

lectotype of Andromeda linearifolia Lesq. USNM 313291 

(pl. 10, fig. 10). 

30 Spermatopslda of Oligocene Creede Flora, Colorado 

Holotype.-VCMG 34078. 

Paratypes.-VCMG 34079, 34080; UCMP 7217, 

7355A, B, 7359, 7379, 7637A, B ( =7638), 7639. 

Genus and Species Indetermlned 

Plate 11, figures 3, 8 

Wolfe and Schorn (1989) designation: Not included. 

Discussion.-Two poorly preserved specimens 

have the same dental morphology: very large, almost 

equal sided, and typically B-3 teeth. One specimen has 

three laminar segments, and the second specimen has 

two laminae that are positioned to indicate a similar 

foliar organization. Because (1) the segments of the first 

specimen are attached to a rachis, (2) the second 

specimen appears to have probably had the same 

organization, and (3) no unattached segments have been 

found, the lamina is interpreted as tripalmatisect (or 

possibly pinnatisect) rather than truly compound (Wolfe 

and Wehr, 1988). In the better preserved specimen, the 

teeth appear to be weakly rosoid. Several extant genera 

of Rosoideae (for example, Potentilla and allied genera) 

have this general leaf architecture, and the fossils may 

represent a tripalmatisect species of Potentilla. Until 

well-preserved specimens are collected, we prefer to 

leave this fossil taxon unnamed. 

Specimens.- UCMG 19766, 34081. 

Subfamily PRUNOIDEAE 

?Genus OSMARONIA Greene 

Osmaronia? stewartlae (Axelrod) Schorn & Wolfe, n. comb. 

Plate 11, figures 4, 9; text figure 1lA 

Arbutus stewartii Axelrod, 1987 (part, typic), p. 146, pl. 34, fig. 1. 

Wolfe and Schorn (1989) designation: Osmaronia? sp. 

Description. -Leaf simple; stipulate?; shape symmetric, narrowly elliptic; base symmetric, acute, apex 

acute and mucronate; length 6.8 em, width 1.8 em; 

venation pinnate; midrib moderate basally, thinning 

markedly and terminating in the apparently glandular 

mucro; 11 pairs of secondary veins, originating at angles 

of 45°-50° (higher apically), decurrent near base of 

lamina, convex, thin, brochidrodromous to eucamptodromous; intersecondary veins composite, thin, extending 

about half the distance to margin; tertiary veins R-R to 

A-0, widely spaced, reticulate; fourth-order veins 

forming imperfect, irregularly polygonal areoles; veinlets 

dendritically branching; external secondary veins three to 

four per secondary vein, originating acutely, forming 

loops of irregular shape; margin entire, erose; petiole 0.8 

em long. 

Discussion.-The two paratypes of Arbutus stewartii are excluded from this species. Both specimens 

lack their original margins and possibly are poorly 

preserved leaves of Populus larsenii. 
F;gure 11. Drawing showing venation of Osmaronia and 

r1odern comparisons. A, 0.? stewartiae (Axelr.) Schorn & 

'''olfe, holotype UCMG 19778 (holotype of Arbutus stewartii 

l'><:elr.). 8, 0. cerasiformis {Torr. & Gray) Greene, extant. C, 

Dichotomanthes tristaniaecarpa Ku rz, extant. D, Arbutus 

rrizonica (Gray) Sarg., extant. 

Leaves of Arbutus arizonica (Gray) Sarg., which 

Fere considered to be closely similar to A. stewartii, have 

closely spaced secondary veins; in a lamina the size of the 

J·olotype of A. stewartii, about 18, rather than 11, pairs of 

~,~condary veins occur. The secondary veins inA. arizonica, although some fork and some are angularly crooked, 

extend approximately straight toward the margin rather 

t., an curve. The angle of origination of secondary veins in 

./1. arizonica is about 60° in the basal half of the lamina 

rather than about 45°. No mucro or indication of glan-

<''ularity occurs on the apex of A. arizonica leaves. 

On the other hand, the same characters that 

<''ifferentiate the holotype of Arbutus stewartii from leaves 

c f Arbutus are found in leaves of Osmaronia. The one 

extant species of the west American Osmaronia, 0. 

cerasifonnis (Torr. & Gray) Greene, has thicker and 

nore strongly brochidodromous secondary veins, 

~~tronger external secondary veins, and a longer mucro 

t., an does A. stewartii. In its thinner secondary veins and 

Peaker loops, the Creede leaf resembles leaves of 

Dichotomanthes and untoothed leaves of Exochorda, 

l: oth extant Chinese genera of Rosaceae. Because only a 

~::ogle specimen of the Creede taxon has been recovered 

rnd because it is not as well preserved as desirable, we 

t"!ntatively and questionably refer it to Osmaronia. 

Citation of a now exclusively Asian genus in the North 

1' merican Tertiary should, we think, be based on better 

material than now available. Had a binomial not been 

based on this specimen, we would not give the specimen 

any formal designation other than "Rosaceae, genus and 

species indetermined." The specimen has sufficient 

characters preserved so that, if better preserved material 

were found at Creede, this material could with 

reasonable certainty be considered conspecific and 

generically determined. 

Nomenclature.-Named after Belle Katherine 

Stewart, the epithet is corrected to stewartiae, m 

accordance with the recommendations of the ICBN. 

Holotype.- UCMG 19778. 

Genus PRUNUS Linnaeus 

Prunus creedensis Axelrod 

Plate 11, figures 2, 5-7; plate 12, figure 5; text figure 124, B 

Prunus creedensis Axelrod, 1987, p. 136, pL 31, fig. 7. 

Sambucus longifolius Axelrod, 1987, p. 150, pL 34, figs. 5, 6. 

Wolfe and Schorn (1989) designation: Prunus creedensis. 

Description. -Leaves simple; stipulate; shape 

symmetric, lanceolate, typically slightly falcate; base 

symmetric, acute to obtuse, apex attenuate; length 

3.8-> 11.5 em, width 1.0-2.6 em; venation pinnate; midrib moderate; 25- > 40 pairs of secondary veins, 

originating at angles of 45°-80°, broadly convex, brochidodromous; numerous composite intersecondary veins, 

some extending to abmedial segment of secondary loop; 

tertiary veins A-0 to typically R-R, percurrent; fourthorder veins forming quadrangular areas; fifth-order veins 

forming with fourth-order veins well-developed, 

polygonal areoles; veinlets branching several times; two 

to four external veins originating from abmedial side of 

each secondary vein, entering teeth medially, craspedodromous; teeth about two to four per secondary vein, 

typically C-1 or D-1; the most basal teeth glandular; 

sinuses angular; petiole 0.6-1.6 em long. 

Discussion.-The holotype of Prunus creedensis has 

a glandular, most basal tooth but no glands on the more 

apical teeth. This character is a strong synapomorphy 

with leaves of extant Prunus. Other characters (for 

example, a closely serrate margin, marked brochidodromy, and craspedodromous external secondary veins) 

are consistent with the assignment to Prunus. 

Three additional leaves of Prunus creedensis are 

represented in the University of Colorado's collections 

and show the holotype to be atypically small. Indeed, 

although the Florissant P. gracilis (Lesq.) MacG. and the 

Troutdale P. treasheri Chan. were differentiated from P. 

creedensis on the basis of their supposedly larger size and 

(or) greater breadth (Axelrod, 1987), these criteria are 

no longer valid. P. creedensis, however, differs from P. 

gracilis in having many more pairs of secondary veins and 

from P. treasheri in having the secondary loops placed 

much closer to the margin. 

Systematic Descriptions and Discussions 31 
The two large laminae that Axelrod (1987) 

referred to Sambucus are here included in Pnmus creedensis. One specimen (Axelrod's pl. 34, fig. 5) has a long 

(1.6 em), distinct petiole and a gland on the most basal 

tooth. Although the second specimen (Axelrod's pl. 34, 

fig. 6, which is the holotype of Sambucus longifolius 

UCMG 19774) lacks a petiole and could thus be 

interpreted as a leaflet, the base of the lamina is actually 

missing. Although we cannot demonstrate whether the 

missing basal teeth were glandular, the intercostal and 

intramarginal venation and dental morphology of this 

specimen are the same as in P. creedensis (fig. 12). 

Prunus creedensis was compared (Axelrod 1987) to 

the extant Asian P. davidiana (Carr.) Franch., but this 

relationship is distant. Whereas P. creedensis has veins 

that enter the teeth medially, P. davidiana has veins that 

extend to the dental sinuses (fig. 12D). The suprabasal 

teeth in P. creedensis are sharp and can have elongated, 

conical glands, but the teeth in P. davidiana are rounded 

and are eglandular or have short, rounded glands. In 

addition, the tertiary veins are typically R-R (some are 

A-0) in P. creedensis but are markedly and uniformly 

A-0 in P. davidiana. Some teeth are entered medially in 

P. pennsylvanica L. f. and most have elongated, conical 

glands, but typically veins extend to the sinuses. Leaves of 

subgenus Padus, however, typically have sharp teeth that 

are entered medially by veins that originate abmedially 

from brochidodromous loops (for example, P. pilosiuscula [Schneid.] Koehne; fig. 12C), and these leaves also 

have marked intersecondary veins and R-R tertiary 

veins. Prunus creedensis appears to represent an extinct 

member of Padus that mimicked the willowlike leaves in 

some other subgenera of Prunus. 

1 mm 

LJ 

Figure 12. Drawing showing venation of Prunus. A, P. 

creedensis Axelr., holotype UCMP 7819. 8, P. creedensis 

Axelr., hypotype UCMG 19774 (holotype of Sambucus /ongifo/ius Axe/r.).C, P. pi/osiuscu/a (Schneid.) Koehne, extant. D, 

P. davidiana (Carr.) Franch., extant. 

32 Spermatopsida of Oligocene Creede Flora, Colorado 

Nomenclature.-Although the type specimens of 

Sambucus longifolius are better preserved and more 

typical in size than the single primary type of Prunus 

creedensis, the epithet creedensis must be accepted as 

valid. 

Holotype.-UCMP 7819. 

Hypotypes.-UCMP 7855-7859; UCMG 19774, 

34082-34084; USNM 422593. 

Prunus sp. 

Pnmus chaneyii auct. non Condit. Axelrod, 1987, p. 135, pl. 28, fig. 4. 

Wolfe and Schorn (1989) designation: Pnmus sp. 

Discussion.-The single Creede leaf referred by 

Axelrod (1987) to Prunus chaneyi validly represents 

Prunus. The most basal teeth on both sides of the lamina 

have glands, an architecture characteristic of this genus. 

In having markedly brochidodromous secondary veins 

and widely spaced, R-R, percurrent tertiary veins, the 

Creede specimen is a member of subgenus Padus, a 

taxon some systematists consider to be of generic rank. 

The Creede leaf is similar in gross characters to leaves of 

P. virginiana L., a similarity also suggested by Axelrod 

(1987). 

Nomenclature.-One cotype of Prunus chaneyi 

(Condit's, 1938, pl. 5, fig. 4) has A-0 tertiary veins and 

salicoid teeth; this specimen is probably an unusually 

small leaf of the Salix that is abundant at many of the 

Neroly localities. The second ootype has an entire margin 

and probably represents Nyssa. The Creede leaf cannot 

be assigned to P. chaneyi, but this single leaf in which 

areolar venation is not preserved is insufficient basis for 

a new species. 

Specimen.-UCMP 7818. 

Family LEGUMINOSAE 

Genus CERCIS Linnaeus 

Cercis sp. 

Text figure 13B 

Cercis buchananensis auct. non Condit. Axelrod, 1987, p. 139, pl. 32, fig. 11. 

Wolfe and Schorn (1989) designation: Cercis sp. 

Discussion.-The top of the petiole of the single 

specimen is swollen and has some vague indications of 

cross striations, thus indicating the presence of the 

0.0\:p~ c 

Figure 13. Drawing showing basal configuration of Cercis. 

A, C. occidenta/isTorr. & Gray. 8, C. sp., UCMG 19775, C, C. 

chinensis Bunge. 
characteristic pulvinus of Leguminosae. In combination 

with simple leaf organization and marked actinodromy, 

assignment to Cercis is probably valid. 

Cercis buchananensis was validly compared by 

Condit (1944) to the extant C. occidentalis Torr. & Gray. 

The leaves of both species have cordate bases that extend 

in a continuously convex curve abmedially and then 

apically: that is, from the top of the petiole to a point 

slightly apical of the petiole the margin describes an 

approximate circle, with the most basal points of the 

cordate base on lines approximately parallel with the 

midrib and approximately half the distance from the 

midrib to the margin (fig. 13). The Creede leaf, however, 

has a margin that is circular only to the most basal points 

of the base and then markedly changes to a much flatter 

curvature; the most basal points of the base on lines 

parallel to the midrib are closer to the midrib than in C. 

occidentalis. This basal configuration is found in C. 

chinensis Bunge (fig. 13C). 

Specimen.-VCMG 19775. 

Genus and Species Indetermined 

Robinia califomica auct. non Axelrod, 1939. Axelrod, 1987 (part), p. 140, pl. 32, 

figs. 1-5. 

Symphoricarpos wassukana auct. non Axelrod, 1956. Axelrod, 1987, p. 149, pl. 33, 

figs. 1, 2. 

Wolfe and Schorn (1989) designation: Legume. 

Discussion.- None of these presumed leaflets is 

well preserved, and, even among the specimens assigned 

by Axelrod (1987) to Robinia, only one shows the crossstriated pulvinus that occurs throughout extant Leguminosae and on even moderately well preserved fossil 

legume foliage. The fossils lack preservation of venation 

above the third order. As preserved, these leaflets could 

be matched by leaflets of a large number of legume 

genera. 

As MacGinitie (1969, 1974) emphasized, even 

well-preserved legume foliage offers major problems in 

identification. Although extensive study of leaf 

architecture in extant and fossil legumes now underway 

by P.F. Herendeen may provide criteria for differentiating well-preserved fossil legume foliage at the 

generic level, the preservation of the Creede fossils is too 

poor for generic or specific determination. 

The two specimens of Symphoricarpos wassukana 

illustrated by Axelrod (1987) are probably short, broad 

variations of the Creede legume leaflet. Although UCMP 

7853 has a very rounded apex, the second specimen 

(UCMP 7851) has an apex no more rounded than one 

legume leaflet (UCMP 7830). Relative to shape of the 

base, one Symphoricarpos specimen (UCMP 7851) has a 

base like one legume specimen (UCMP 7833), and the 

second Symphoricarpos specimen (UCMP 7853) has a 

base like a second legume specimen (UCMP 7835). One 

unfigured cited hypotype of S. wassukana (UCMG 

19784) was also cited as a paratype of Quercus creedensis 

(Axelrod, 1987, p. 108). 

Specimens.-UCMP 7829, 7830, 7832-7835, 7851, 

7853, 7861. 

Family BIGNONIACEAE 

Genus CATALPA Scopoli 

Catalpa coloradensis Schorn & Wolfe, n. comb. 

Plate 12, figures 1, 2, 7 

Nuphar coloradensis Axelrod, 1987, p. 108, pl. 33, figs. 4, 5. 

Wolfe and Schorn (1989) designation: Catalpa sp. 

Description. -Leaves simple; shape symmetric, 

widely to very widely ovate; base symmetric, cordate, 

apex unknown; length unknown but > 10 em, width 

10-12 em; venation perfectly actinodromous; nine primary veins; five primary veins including midrib massive; 

most apical pair of lateral primary veins convex, 

extending apically to beyond the same distance as the 

origination point of the second most basal pair of medial 

secondary veins, typically slightly convergent with medial 

secondary veins, brochidodromous or, if lateral lobes 

present, craspedodromous; four to five secondary veins 

originating at angles of 30°-40° from basal sides of most 

apical pair of lateral primary veins, convex, brochidrodromous; second most apical pair of lateral primary veins 

extending apically to the same (or less) the distance as 

the origination point of the most basal pair of medial 

secondary veins, slightly convergent with secondary veins 

of suprajacent lateral primary vein, convex, brochidodromous; three to five pairs of secondary veins originating at 

angles of 30°-40° from basal sides of second most apical 

pair of lateral primary veins, convex, brochidrodromous; 

third most apical pair of lateral primary veins extending 

abmedially and away from apex, slightly convergent with 

secondary veins of suprajacent lateral primary vein, 

convex, brochidodromous; two to four pairs of secondary 

veins originating acutely from basal sides of third most 

apical pair of lateral primary veins, convex, brochidodromous; most basal pair of lateral primary veins about 

same thickness as seC9ndary veins of suprajacent lateral 

primary vein, brochidodromous; tertiary veins R-R, percurrent, some branching; higher order venation not 

preserved; brochidodromous loops formed by an 

external vein of suprajacent secondary vein that joins 

subjacent secondary vein; abmedial side of loop extends 

almost to margin; some external veins originating acutely 

from apical sides of most apical pair of lateral primary 

veins, brochidodromous; margin entire, thickened but no 

fimbria! vein; some leaves have a single lateral lobe that 

has a rounded apex; petiole very thick, > 7 em long, with 

broad base. 

Discussion.-The distinctive leaves of Catalpa coloradensis are highly dissimilar to leaves of Nuphar. 

Leaves of Nuphar (1) have an auriculate base, (2) are 

basically pinnately veined, (3) have major veins that 

markedly diverge from one another, ( 4) have secondary 

Systematic Descriptions and Discussions 33 
veins that fork into branches of approximately equal size 

(that is, dichotomize) two or three times, with the first 

for king typically occurring about half the distance (or 

less) to the margin, and (5) are unlobed. The broad 

petiolar base (pl. 12, fig. 2) represents the presumed 

plane of dehiscence. The leaves of Nymphaeaceae die 

attached to plant and are dissociated by degradation of 

the petiole. The specimen (UCMP 7523) considered to 

be a nymphaeaceous "rootscar" (Axelrod, 1987, p. 17) is 

a poorly preserved impression of unknown affinities. 

The specimen illustrated as figure 2 on plate 12 is 

the most complete lamina of Catalpa coloradensis known, 

although the preserved margin has apparently resulted 

from degradation of the original margin. One paratype 

(pl. 12, fig. 7), although a fragment, illustrates the margin 

and adjacent venation. The holotype (Axelrod, 1987, pl. 

22, fig. 5) is perhaps lobed, but the margin is sufficiently 

decayed to be uncertain about lobing; one paratype (pl. 

12, fig. 1), however, is unquestionably lobed. 

Deeply cordate leaves that are actinodromous, 

have as many as nine primary veins, have an entire 

margin that can be lobed but that does not have a 

fimbrial vein, and have secondary veins that are markedly 

brochidodromous close to the margin, occur primarily in 

the palmate dilleniids (Hickey and Wolfe, 1975). In the 

palmate dilleniids, however, the venation around the top 

of the petiole tends to have a spiderweb appearance: the 

tertiary veins are arranged concentrically around the top 

of the petiole, and the fourth-order venation is strong and 

orthogonal to the tertiary veins. On the other hand, the 

Creede leaves are very similar in primary, secondary, and 

tertiary venation patterns to leaves of Catalpa (Bignoniaceae) and Paulownia (Scrophulariaceae ), which are 

considered to be closely related to one another (Airy 

Shaw, 1985). Catalpa especially can have leaves that (1) 

have a thick petiole and major veins, (2) can be lobed, 

and (3) have markedly brochidodromous secondary 

veins. Lobes that have rounded apices are found in leaves 

of the Asian C. ovata D. Don. 

Holotype.-VCMP 7520. 

Paratypes.-VCMP 7518, 7519, 7521, 7522. 

Hypotype.-UCMG 34085. 

LILIATAE, Family lndetermlned 

Genus MONOC01YLOPHYLLUM Chandler 

Monocotylophyllum sp. 

Cyperacites creedensis Axelrod, 1987, p. 100, pl. 17, fig. 1. 

Discussion. -In order for leaf impressionscompressions to be placed in Cyperaceae, they should 

have evidence of a triangular cross section. In fact, the 

Creede leaves assigned to Cyperacites are described as 

"grass-like." The morphology of many monocotyledonous leaves, including these Creede specimens, is too 

34 Spermatopsida of Oligocene Creede Flora, Colorado 

simple to determine even the family represented. No 

point is served by giving an epithet to such material. 

Specimen.-UCMP 7495. 

Names of Dubious Value 

Species, as well as citations of previously described 

species, based on fragmentary and (or) poorly pre.served 

specimens create nomenclatorial as well as taxonomic 

problems. Although the ICBN allows for formal 

rejection of such names, the procedure is long. In the 

previous discussions of systematics of the Creede 

conifers, we suggested that some formally named species 

were based on material that is too fragmentary or poorly 

preserved for valid characterization of the species: Picea 

coloradensis Axelr. and Tsuga petranensis Axelr. 

Following we list and discuss other species based on 

Creede material that we consider also to be good 

candidates for rejection; citations based on Creede 

specimens of species previously described from other 

floras are also listed and discussed, except for those 

citations previously included in our formal synonymies. 

Quercus creedensis Axelrod, 1987, p.107, pl. 21, figs. 2-5. 

PL'lte 12, figures 4, 6 

Discussion.-These specimens are all entiremargined. Some indications of secondary veins are 

present, but not even the manner of looping can be 

determined (pl. 12, figs. 4, 5). Tertiary and higher-order 

venation is totally lacking; the basis for Axelrod's (1987, 

p. 107) description of tertiary and higher order venation 

is unknown. The shortness of the petiole suggests that it 

could be a petiolule; concomitant with the asymmetry of 

the laminae, these laminae may represent very poorly 

preserved specimens of the indetermined legume. 

Specimens.-VCMP 7514-7517; UCMG 19784 

(note that this cited paratype was also cited (Axelrod, 

1987, p. 149) as a hypotype of Symphoricarpos wassukana). 

Hippurus coloradensis Axelrod, 1987, p. 143, pl. 33, fig. 9. 

Discussion.-The specimen on which Hippwus col-~ 

oradensis is based is an axis that has three nodes, to which 

three smaller axes are attached. One node has a 

filamentous structure also attached. The three smaller 

axes also have nodes, to which smaller filamentous 

structures are attached. Striations on the axes suggest 

some vascularization. The specimen is not diagnostic as 

to whether it is even an angiosperm; included in the 

University of Colorado's Creede collections is a similar 

specimen (UCMG 19689) that has the nodal sheaths of 

Equisetum. 

Specimen.- UCMP 9840. 
Shepherdia creedensis Axelrod, 1987, p. 145, pl. 34, fig. 3. 

Discussion.-The holotype of Shepherdia creedensis lacks preservation of the original margin and of 

venation above the tertiary level. The broadly ovate 

shape, truncate base, irregularly spaced secondary veins, 

and thin, undulatory tertiary veins are characters that are 

found also in Populus larsenii. This may be a poorly 

preserved specimen of P.larsenii, but without the margin 

we cannot be certain. 

Specimens.- UCMP 7844, 7845. 

Chilopsis coloradensis Axelrod, 1987, p. 150, pl. 34, fig. 7. 

PL'lte 12, figure 3 

Discussion.-Although the leaf of Chilopsis coloradensis is linear, the photographic enlargement (pl. 12, 

fig. 3) indicates much compression by folding. What 

superficially appears to be strongly decurrent secondary 

veins that have a very low angle of origination are folds in 

the lamina. The margin appears to be entire, but folding 

makes this impossible to determine certainly. The 

specimen is possibly a folded lamina of Eleiosina praeconcinna. 

Specimen.-UCMP 7852. 

Fraxinus creedensis Axelrod, 1987, p. 148, pl. 33, fig. 6. 

Plate 12, figure 8 

Discussion.-As shown in the photographic 

enlargement (pl. 12, fig. 7), the "leaflets" of this specimen 

have, where not degraded, sharp B-2, closely spaced 

serrations. The secondary veins originate at angles of 

about 35° and are brochidodromous; the teeth are 

entered by external secondary veins, which appear to 

enter near the apical side of the tooth. 

The preserved characters of Fraxinus creedensis 

can be found in several families of Northern Hemisphere 

Rosidae (as redefined by Wolfe, 1989): Juglandaceae, 

Rosaceae, Araliaceae, and Oleaceae (which includes 

Fraxinus). An additional, poorly preserved specimen in 

the University of Colorado's collections appears also to 

represent F. creedensis and has five "leaflets" attached to 

a rachis. The fact that both specimens have laminae still 

attached to a rachis suggests that the leaf was not truly 

compound but was pinnatisect (Wolfe and Wehr, 1988), 

a condition that commonly occurs in Rosaceae and 

Sambucaceae. The poor preservation of both specimens 

prohibits further speculation on affinities. 

Specimen.-VCMG 19777. 

Picea lahontensis auct. non MacGinitle. Axelrod, 1987 (part), p. 85. 

Discussion.-A defoliated axis (UCMP 7295) 

appears to have irregularly arranged bud and leaf scars 

and probably represents an angiosperm. A supposed 

winged seed (UCMP 7301) could be either a floral organ 

or a fragment of a monocot leaf; the "seed" area of the 

specimen is an artifact of wrinkling, and the specimen has 

weak parallel venation (accompanied by transverse 

veins) the entire length. 

Specimens.- UCMP 7295, 7301. 

Pseudotsuga glaucoides Axelrod, 1987 (part, nontyplc), p. 96, pl.lS, 

fig. 4. 

Discussion.-This supposed ovulate cone scale 

lacks a bract scale, impression of the seeds or wings or 

any other character that is diagnostic of a pinaceous cone 

scale. What taxon or organ this specimen represents is 

problematic. 

Specimen.- UCMP 7425. 

Sapindus coloradensis auct. non Cockerell. Axelrod, 1987, p.142, pl. 

33, fig. 7. 

Discussion.-The falcate shape and attenuate apex 

would appear to differentiate this specimen from the 

legume taxon called Robinia califomica (our "legume"): 

however, one "Robinia," UCMP 7833, is also somewhat 

falcate and has an acute apex. UCMP 7833 also has an 

asymmetrically, broadly rounded base, as does the 

"Sapindus." A second "Robinia," UCMP 7829, does not 

have a complete apex, but the margin changes to a 

concave curvature, which suggests that an attenuate apex 

was present. In the "Sapindus," however, the secondary 

veins are "poorly preserved" and "finer nervation [is] not 

preserved" (Axelrod, 1987, p. 142). Further discussion is 

pointless. 

Specimen.-UCMP 7899. 

Condalia mohavensis auct. non Axelrod, 1939. Axelrod, 1987, p. 144, 

pl. 33, fig. 8. 

Discussion.-The specimen is partially covered by 

a mineral growth (stain), and details of venation and 

character of the margin are thus uncertain. The specimen 

has suprabasal secondary veins that are "crowded" at the 

base as in Populus, and two conspicuous, rounded stains 

are present on either side of the top of the petiole; these 

stains could represent the acropetiolar glands of Populus. 

Specimen.-UCMP 7841. 

Zizyphus florissantii auct. non (Lesquereux) MacGinitie. Axelrod, 

1987' p. 144, pl. 34, fig. 4. 

Discussion.-In Zizyphus florissantii, the lateral 

major veins are close to the margin and are connected to 

the midrib by numerous percurrent veins (MacGinitie, 

1953). In this Creede specimen, however, (1) the lateral 

primary veins have external veins that form elongated 

brochidrodromous loops between the margin and the 

lateral primary veins, and (2) strong intersecondary veins 

extend apically between the lateral primary veins and the 

midrib, and thus no percurrent veins are present. This 

Creede specimen cannot represent Z. florissantii. In 

these two characters, concomitant with the actinodromous three primary veins and the linear shape, this 

Names of Dubious Value 35 
specimen is similar to Berberis coloradensis and may be 

nothing more than a large, battered lamina of that 

species. 

Specimen.-VCMP 7843. 

Summary of Systematic Revisions 

Of the 42 new specific epithets based on Creede 

material by Axelrod (1956, 1987), we recommend 

rejection of 10 because the basionyms are poorly 

preserved and (or) fragmentary and either cannot be 

distinguished from previously named species of the same 

genera or lack characters sufficient for familial 

determination. Eight of Axelrod's (1987) new species 

and one new combination we accept as valid and properly 

assigned to their respective genera. Of the remaining 

species, 5 are transferred to other genera and 19 are 

synonymized under other species. The following list 

summarizes our recommended disposition of these 

epithets. 

Picea coloradensis Axelr.: Recommend rejection. 

Pinus engelmannoides Axelr.: Recommend rejection. 

P. ponderosoides Axelr.: Synonymized under P. crossii Knowlt. 

P. riogrande Axelr.: Accepted. 

P. sanjuanensis Axelr.: Accepted. 

P. wasonii Axelr.: Synonymized under P. crossii Knowlt. 

Tsuga petranensis Axelr.: Recommend rejection. 

Juniperus creedensis Axelr.: Accepted. 

J. gracillensis Axelr.: Synonymized under Eleopoldia lipmanii (Axelr.) 

Schorn & Wolfe. 

Cyperacites creedensis Axelr.: Recommend rejection. 

Populus creedensis Axelr.: Synonymized under Populus larsenii 

(Knowlt.) Schorn & Wolfe. 

Salix creedensis Axelr.: Synonymized under Eleiosina praeconcinna 

(Cocker.) Schorn & Wolfe. 

Quercus creedensis Axelr.: Recommend rejection. 

Nuphar coloradensis Axelr.: New combination as Catalpa coloradensis 

(Axelr.) Schorn & Wolfe. 

Ranunculus creedensis Axelr.: New combination as Potentilla creedensis 

(Axelr.) Schorn & Wolfe. 

Berberis coloradensis Axelr.: Accepted. 

B. riogrande Axelr.: Synonymized under B. coloradensis Axelr. 

Mahonia creedensis Axelr. [nom. nud.]: Synonymized under Mahonia 

aceroides (Knowlt.) Schorn & Wolfe. 

Ribes birdseyii Axelr.: Synonymized under R lacustroides Axelr. 

R. creedensis Axelr.: Synonymized under R lacustroides Axelr. 

R. dissecta Axelr.: Synonymized under R lacustroides Axelr. 

R. lacustroides Axelr.: Accepted. 

R riogrande Axelr.: Synonymized under R lacustroides Axelr. 

R. stevenii Axelr.: An automatic junior synonym of R. riogrande Axelr., 

which is synonymized under R lacustroides Axelr. 

R. wasonii Axelr.: Synonymized under R /acustroides Axelr. 

Fend/era coloradensis Axelr.: Synonymized under Berberis coloradensis 

Axelr. 

Jamesia caplanii Axelr.: Accepted. 

Philadelphus creedensis Axelr.: Synonymized under Berberis coloradensis Axelr. 

Crataegus creedensis Axelr.: Accepted. 

Fallugia lipmanii Axelr.: New combination as Eleopoldia lipmanii 

(Axelr.) Schorn & Wolfe. 

36 Spermatopsida of Oligocene Creede Flora, Colorado 

Physocarpus petiolaris Axelr.: Transferred to Ribes as R. robinsonii 

Schorn & Wolfe (a new epithet required because of homonomy). 

P. triloba Axelr.: Synonymized under Ribes robinsonii Schorn & Wolfe. 

(triloba would also be a homonym in Ribes). 

Prnnus creedensis Axelr.: Accepted. 

Rubus riogrande Axelr.: Synonymized under Ribes lacustroides Axelr. 

Sorbus potentilloides (Knowlt.) Axelr.: Accepted. 

Acer riogrande Axelr.: Synonymized under Ribes lacustroides Axelr. 

Hippurus coloradensis Axelr.: Recommend rejection. 

Shepherdia creedensis Axelr.: Recommend rejection. 

Arbutus stewartii Axelr.: New combination as Osmaronia? stewartiae 

(Axelr.) Schorn & Wolfe. 

Vaccinium creedensis Axelr.: Synonymized under Berberis coloradensis 

Axelr. 

Fraxinus creedensis Axelr.: Recommend rejection. 

Chilopsis coloradensis Axelr.: Recommend rejection. 

Sambucus longifolius Axelr.: Synonymized under Prnnus creedensis 

Axelr. 

Of the 73 species in Axelrod's (1987) Creede flora, 

30 represent citations of species previously described 

from other floras; none of these citations are considered 

valid. Further, many of the nonbasionymic specimens 

assigned to species based on Creede material represent 

other taxa. The following list summarizes the systematic 

disposition of Axelrod's (1987) Creede flora as made in 

our current report. 

Abies conco/oroides R.W. Br.: A. rigida Knowlt. 

A. rigida Knowlt.: A. rigida Knowlt. 

Picea coloradensis Axelr.: Pinus cf. P. crossii Knowlt., Pinus spp. indet., 

Pinaceae gen. indet. 

P. lahontensis MacG.: Abies rigida Knowlt., Picea sp., Pinus cf. P. crossii 

Knowlt., P. spp. indet., Pinaceae gen. indet., Magnoliophyta gen. 

indet. 

P. sonomensis Axelr.: Pinus spp. indet. 

Pinus alvordensis Axelr.: P. sanjuanensis Axelr., P. crossii Knowlt., P. 

spp. indet. 

P. coloradensis Knowlt.: P. crossii Knowlt., P. spp. indet., Pinaceae gen. 

indet. 

P. crossii Knowlt.: P. crossii Knowlt., Pinaceae gen. indet. 

P. engelmannoides Axelr.: P. cf. P. crossii Knowlt. 

P. florissantii Lesq.: P. crossii Knowlt. 

P. macginitiei Axelr.: P. sp. 2. 

P. ponderosoides Axelr.: P.crossii Knowlt., P. cf. P. crossii Knowlt. 

P. riogrande Axelr.: P. cf. P. crossii Knowlt., P. riogrande Axelr., P. sp. 1, 

P. spp. indet. 

P. sanjuanensis Axelr.: P. sanjuanensis Axelr., P. riogrande Axelr. 

P. wasonii Axelr.: P. crossii Knowlt. 

Pseudotsuga glaucoides Axelr.: Abies rigida Knowlt., ?Pinus crossii 

Knowlt., P. spp. indet., Pinaceae gen. indet., indet. plant. 

Tsuga petranensis Axelr.: Pinus spp. indet., Pinaceae gen. indet. 

Juniperus creedensis Axelr.: J. creedensis Axelr. 

J. gracillensis Axelr.: Eleopoldia lipmanii (Axelr.) Schorn & Wolfe. 

Cyperacites creedensis Axelr.: Monocotylophyllum sp. 

Populus cedrusensis Wolfe: P. larsenii (Knowlt.) Schorn & Wolfe. 

P. creedensis Axelr.: P. /arsenii (Knowlt.) Schorn & Wolfe. 

P. pliotremuloides Axelr.: P. larsenii (Knowlt.) Schorn & Wolfe. 

Salix creedensis Axelr.: Eleiosina praeconcinna (Cocker.) Schorn & 

Wolfe. 

S. venosiuscula H. V. Sm.: S. sp. 

Betula smithiana (Axelr.) Axelr.: Populus /arsenii (Knowlt.) Schorn & 

Wolfe. 

Quercus creedensis Axelr.: Magnoliatae indet. (possibly Leguminosae 

gen. indet. ). 
Nuphar coloradensis Axelr.: Catalpa coloradensis (Axelr.) Schorn & 

Wolfe. 

Ranunculus creedensis Axelr.: Potentilla creedensis (Axelr.) Schorn & 

Wolfe. 

Berberis coloradensis Axelr.: B. coloradensis Axelr. 

B. riogrande Axelr.: B. coloradensis Axelr. 

Mahonia creedensis Axelr.: M. aceroides (Knowlt.) Schorn & Wolfe. 

M. obliqua MacG.: M. aceroides (Knowlt.) Schorn & Wolfe. 

Ribes birdseyii Axelr.: R lacustroides Axelr. 

R. creedensis Axelr.: R lacustroides Axelr. 

R. dissecta Axelr.: R lacustroides Axelr. 

R. lacustroides Axelr.: R lacustroides Axelr. 

R. riogrande Axelr.: R lacustroides Axelr. 

R. stevenii Axelr.: R lacustroides Axelr. 

R. wasonii Axelr.: R lacustroides Axelr. 

R. webbii Wolfe.: R lacustroides Axelr. 

Fend/era coloradensis Axelr.: Berberis coloradensis Axelr. 

Jamesia caplanii Axelr.: J. caplanii Axelr. 

Philadelphus creedensis Axelr.: Berberis coloradensis Axelr. 

Cercocarpus holmesii (Lesq.) Axelr.: Eleiosina praeconcinna (Cocker.) 

Schorn & Wolfe, C. henricksonii Schorn & Wolfe. 

C. linearifolius (Lesq.) Axelr.: C. nanophyllus Schorn & Wolfe. 

Chamaebatiaria creedensis R.W. Br.: Stockeya creedensis (R.W. Br.) 

Wolfe & Wehr. 

Crataegus creedensis Axelr.: C. creedensis Axelr. 

Fallugia lipmanii Axelr.: Eleopoldia lipmanii (Axelr.) Schorn & Wolfe. 

Holodiscus hameyensis (Am.) Axelr.: Ribes obovatum Schorn & Wolfe, 

Holodiscus stevenii Schorn & Wolfe. 

H. idahoensis Chan. & Axelr. [nom. nud.]: H. stevenii Schorn & Wolfe. 

Peraphyllum septentrionalis (Lesq.) Axelr.: Eleiosina praeconcinna 

(Cocker.) Schorn & Wolfe. 

Physocarpus petiolaris Axelr.: Ribes robinsonii Schorn & Wolfe. 

P. triloba Axelr.: Ribes robinsonii Schorn & Wolfe. 

Pnmus chaneyii Cond.: P. sp. 

P. creedensis Axelr.: P. creedensis Axelr. 

Rosa hilliae Lesq.: Holodiscus stevenii Schorn & Wolfe. 

Rubus riogrande Axelr.: Ribes lacustroides Axelr. 

Sorbus potentilloides (Knowlt.) Axelr.: S. potentilloides (Knowlt.) Axelr. 

Cercis buchananensis Cond.: C. sp. 

Robinia califomica Axelr.: Leguminosae gen. indet. 

Acer riogrande Axelr.: Ribes lacustroides Axelr. 

Sapindus coloradensis Cocker.: Magnoliatae gen. indet. (possibly Leguminosae gen. indet. ). 

Hippurus coloradensis Axelr.: indet. plant (possibly Equisetwn). 

Condalia mohavensis Axelr.: Magnoliatae gen. indet. (possibly Populus 

larsenii [Knowlt.] Schorn & Wolfe. 

Zizyphus florissantii (Lesq.) MacG.: Magnoliatae gen. indet. (possibly 

Berberis coloradensis Axelr.). 

Shepherdia creedensis Axelr.: Magnoliatae gen. indet. (possibly Populus 

larsenii (Knowlt.) Schorn & Wolfe). 

Arbutus stewanii Axelr.: Osmaronia? stewaniae (Axelr.) Schorn and 

Wolfe, Magnoliatae gen. indet. (possibly Populus larsenii [Knowlt.] 

Schorn & Wolfe). 

Vaccinium creedensis Axelr.: Berberis coloradensis, Magnoliatae gen. 

indet. 

Fraxinus creedensis Axelr.: Magnoliatae gen. indet. 

Symphoricarpos wassukana Axelr.: Leguminosae gen. indet. 

Chilopsis coloradensis Axelr.: Magnoliatae gen. indet. (possibly Eleiosina praeconcinna [Cocker.] Schorn & Wolfe). 

Sambucus longifolius Axelr.: Prnnus creedensis Axelr. 

DESCRIPTION J'ND DISCUSSION OF A 

NEOGENE SPRUCE 

Family PINACEAE 

Genus PICEA Dietrich 

Picea lahontensis MacGinitie 

Picea lahontense MacGinitie, 1933 (part, typic) p. 46, pl. 3, figs. 6, 8 only. 

Pseudotsuga masoni MacGinitie, 1933, p. 47, pl. 3, figs. 1-3. Oliver, 1934, p. 14. 

Dorf, 1938, p. 107, pl. 1, fig. 3. 

Pinus hameyana Chaney and Axelrod, 1959 (part, typic), p. 141, pl. 13, figs. 1, 2. 

Axelrod, 1964 (part), p. 108, pl. 6, figs. 20-22. 

Picea magna auct. non MacGinitie. Axelrod, 1956, p. 275, pl. 4, figs. 7-12; pl. 25, 

figs. 8, 9. Chaney and Axelrod, 1959, p. 140, pl. 12, figs. 10-15. Axelrod, 

1964, p. 108, pl. 6, figs. 9-13. Wolfe, 1964, p. 15, pl.1, figs. 3, 5; pl. 6, figs. 7, 

12, 17, 18, 22. Graham, 1%5, p. 58, pl. 3, figs. 9, 10. 

Pseudotsuga sonomensis auct. non Dorf. Axelrod, 1956 (part), p. 277, pl. 4, fig. 21. 

Picea breweriana auct. nonS. Watson. Wolfe, 1964 (part), p. 14, pl. 6, figs. 4, 5, 

8, 9. 

Pinus alvordensis auct. non Axelrod, 1944. Axelrod, 1985, p. 114, pl. 17, figs. 

15-19. 

Conifer of unknown affinities. Chaney and Axelrod, 1959, p. 144, pl. 14, fig. 9. 

Emended description.- Ovulate cone incomplete 

(small part of apex missing); shape lanceolate, slightly 

asymmetrical, base obtuse; 11.8 em long, 3.5 em wide; 

stalk 0.2 em long; cone scales as wide as 1.5 em, 0.9 em 

high, width to height ratio about 1.6:1, apex rounded and 

with a small notch, margin smooth to incipiently crenulate, weak vertical ribbing, scales appearing woody, no 

exposed bract scale. Winged seed has cellular pattern 

parallel with long axis of wing; wing 1.2-1.4 em long, 

0.7-1.0 em wide, wing-length to wing-width ratio 1.6:1; 

seed surface convex, indentation at proximal end on 

admedial edge, proximal end truncate, 0.7 em long, 

0.4-0.5 em wide, seed-length to seed-width ratio 1.5:1; 

length of seed and wing 1.8-2.1 em, wing-length to 

seed-length ratio 2:1. Foliated axis more than 3.8 em 

long, 0.4 em wide in apical part, furnished with pulvinae 

about 0.2 em long that are directed about 70° from 

vertical plane of main axis; leaf scar shape on adaxial 

surface rounded and on abaxial surface triangular; leaves 

at least 3.2 em long, 0.2 em wide, base tapers slightly at 

base, distinct keel on abaxial side. 

Discussion.-When MacGinitie (1933) described 

Picea lahontensis (asP. /ahontense), only one other fossil 

specimen from conterminous United States had been 

referred to the genus. Considerable confusion now exists 

regarding the circumscription of MacGinitie's species, 

which is related to the Creede spruce. 

The holotype of Picea /ahontensis (MacGinitie, 

1933, pl. 3, fig. 8) is a partially foliated axis that has four 

attached leaves. The figure was retouched and shows the 

lower righthand leaf as complete. Further preparation 

shows, however, that the apex of this leaf is missing (as 

are the apices of the other leaves); however, tapering at 

the point of the break indicates that the leaf is almost 

complete. Additional material from the Trout Creek 

locality and deposited in UCMP has complete leaves, 

Description and Discussion of a Neogene Spruce 37 
which are, as leaves of the holotype, broad for Picea 

(about 1 mm). The epithet lahontensis, which is well 

entrenched in the literature, can thus be retained by 

basing an emended description on topotypic material that 

includes complete leaves, cones, and winged seeds that 

were originally referred to Pseudotsuga masoni. 

Three morphotypes of foliated axes of Picea occur 

in th: Neogene of western. North America: one morphotype Is represented by the specimen from a Mascall flora 

locality originally assigned to Abies chaneyi Mason 

(Mason, 1927) and later transferred by Axelrod (1976) to 

Cephalotaxus bonseri (Knowlt.) Chan. & Axelr.; a second 

morphotype is represented by the Tipton specimen that 

':as first referred to A. chaneyi by Brown (1937; pl. 45, 

fig. 24) and later transferred by him (Brown, 1940) to A. 

concoloroides R.W. Br.; and a third morphotype is 

represented by P. lahontensis, which occurs at Tipton as 

well as Trout Creek. All three morphotypes have the 

pulvinae (peglike structures on the axis) and leaves that 

are unconstricted or only slightly constricted and thus do 

not form a petiole; this leaf base is, in Pinaceae, restricted 

to Picea leaves. 

The defoliated axis from Tipton that Mason (1927 

pl. 3, fig. 2) assigned to Picea sp. (Chaney and Axelrod' 

19~9, reassigned the specimen toP. sonomensis Axelr.) i~ Abzes. Structures that appear to be the pulvinae 

characteristic of Pice a are the persistent bud scales of the 

male cones characteristic of Abies and Keteleeria as are 

the circular leaf scars of the fossil. Whereas K;teleeria 

typically has two or three male cones per bud, the fossil 

and extant Abies have one male cone per bud. 

. In the absence of organic connection, the 

assign~ent of the large cone (MacGinitie, 1933, pl. 3, fig. 

6) to Pzcea lahontensis is uncertain. This Picea cone and 

the P. lahontensis foliage are, however, the only morpho types of their respective organs of Picea found at 

Trout Creek, and both also occur at Tipton (the second 

morphotype of Picea foliage also occurs at Tipton). We 

tentatively consider the large cone to belong to P. lahontensis. The large Picea winged seeds that formed the basis 

for Pseudotsuga masoni MacG. (MacGinitie, 1933) 

would readily be accommodated on the large cone scales of 

this cone type, and similar large cones occur in 

association with large winged seeds at both Tipton and 

Hidden Lake. 

. . The . winged seed (MacGinitie, pl. 3, fig. 4) 

ongmally mcluded as a paratype of Picea lahontensis 

a~parently has a seed that is not fully developed; of the 7 5 

wmged seeds of Picea known from Trout Creek all 

except this single specimen have large seeds such a; the 

spec~mens. a~si~ned to Pseudotsuga masoni. This single 

specimen IS similar to seeds borne on basal or apical cone 

sc.ales, where typically an ovule is absent and only the 

~mg develops or an ovule aborts early in development; as 

m aborted extant seeds, the Trout Creek specimen 

38 Spermatopsida of Oligocene Creede Flora, Colorado 

represents a flattened, linear seed that is flanked on 

either side by wing tissue that is as wide as the seed. Most 

winged seeds referred to Picea lahontensis by subsequent 

authors represent broken or similarly aborted specimens. 

Winged seeds of Picea sonomensis Axelr. consistently 

differ in having a seed outline that is elongated (length: 

width > 2:1) versus a seed outline that is wide (length: 

width < 2:1) in winged seeds of P. lahontensis. 

Affinities of Picea lahontensis to extant species are 

uncertain. MacGinitie (1933) considered P. lahontensis 

to be similar toP. engelmannii Parry ex Engelm., but this 

comparison was partly based on the atypically small 

winged seed. Axelrod (1985) considered P. lahontensis to 

be related to some Asian spruces, for example, P. polita 

(Sieb. & Zucc.) Carr.; these spruces have angular foursided leaves and thus are very dissimilar. The broad, 

flattened leaves that have a keel only on the abaxial side 

of P. lahontensis suggest that the fossil is an extinct 

member of section Omarika. 

The foliated axis of the conifer of unknown 

affinities (Chaney and Axelrod, 1959) from the Tipton 

flora has the stout pulvinae characteristic of Picea. The 

width of the leaves relative to their length is similar toP. 

lahontensis. The supposed cone of the Tipton specimen 

has features indicative of a spruce gall (H.I. Scudder, oral 

commun., 1982), such as formed on extant species of 

P_ic~a by the spruce gall aphid Chennes ( =Adelges). A 

similar gall was thought to represent an aroid spadix 

(MacGinitie, pl. 7, fig. 4), and an unfigured specimen is 

present in collections of the Stinking Water flora (UCMP 

loc. P4120). 

Holotype.- UCMP 550. 

Paratypes.-VCMP 551, 552. 

Hypotypes.- UCMP 553, 554. 

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Oxford, England, Clarendron Press, p. 75-104. 

Wolfe, J.A., and Schorn, H.E., 1989, Paleoecologic, paleoclimatic, and evolutionary significance of the Oligocene 

Creede flora, Colorado: Paleobiology, v. 15, p. 180-198. 

Wolfe, J.A., and Tanai, Toshimasa, 1987, Systematics, phylogeny, and distribution of Acer (maples) in the Cenozoic 

of western North America: Hokkaido University Faculty 

of Science Journal, ser. 4 (Geology), v. 22, p. 1-246. 

Wolfe, J.A., and Wehr, Wesley, 1987, Middle Eocene 

dicotyledonous plants from Republic, northeastern 

Washington: U.S. Geological Survey Bulletin 1597,25 p. 

___ 1988, Rosaceous Chamaebatiaria-like foliage from the 

Paleogene of western North America: Aliso, v. 12, 

p. 177-200. 
PLATES 1-13 

Contact photographs of the plates in this report are available, at cost, 

from the U.S. Geological Survey Photographic Library, Federal Center, 

Denver, CO 80225 
PLATE 1 

[All figures enlarged] 

FIGURE 1, 2. Juniperns creedensis Axelr. (p. 4). 

1. Paratype UCMP 7452, x2.5. 

2. Paratype UCMP 7460, x 3.5. 

3-10. Abies rigida Knowlt. (p. 9). 

3. Hypotype UCMP 7258, x 2.5. 

4. Hypotype UCMP 7280, x 2.5. 

5. Hypotype UCMP 7279, x 2.5. 

6. Homeotype UCMP 7260, x 2.5. 

7. Hypotype UCMP 7251, x 2.5. 

8. Hypotype UCMP 7410 (homeotype of Pseudotsuga glaucoides 

Axelr.), x 3.5 (note circular leaf scars at top). 

9. Hypotype UCMP 7275, x 9 (lined area is pterostegium). 

10. Homeotype UCMP 7267, x 3.5. 

11, 12. Pinus sanjuanensis Axelr. (p. 11 ). 

11. Paratype UCMP 7239, x 2. 

12. Paratype UCMP 7242, x 2. 

13, 14. Pinus spp., atypical winged seeds (p. 15). 

13. Specimen UCMP 7229 (homeotype of P. alvordensis Axelr.), x 2. 

14. Specimen UCMP 7230 (homeotype of P. alvordensis Axelr.), x 2. 
U.S. GEOLOGICAL SURVEY BULLETIN 1923 PLATE 1 

3 

8 13 14 

JUNIPERUS, ABIES AND PINUS 
PLATE 2 

(All figures natural size unless otheiWise indicated] 

FIGURES 1-9. Pinaceae, ovulate cones (p. 5). 

Arbitrary abrasion stages of ovulate cones of Pinaceae from least abrasion 

(stage 1) to high abrasion (stage 6). 

1. Stage 1, Pinus crossii Knowlt., hypotype UCMP 7354 (p. 12). 

2. Stage 2, P. crossii Knowlt., hypotype UCMP 7216 (hypotype of P. 

florissantii Lesq.) (p. 12). 

3. Stage 3, Pinus cf. P. crossii Knowlt., specimen UCMP 7316 (hypotype 

of Picea lahontensis MacG.) (p. 14). 

4. Stage 4, P. cf. P. crossii Knowlt., specimen UCMP 7353 (paratype of P. 

ponderosoides Axelr.) (p. 15). 

5. Stage 5, Pinaceae, gen. indet., specimen UCMP 7287 (holotype of 

Picea coloradensis Axelr.) (p. 15). 

6. Stage 6, Pinaceae gen. indet., specimen UCMP 7290 (paratype ofPicea 

coloradensis Axelr.) (p. 15). 

7. Stage 6, Pinaceae gen. indet., specimen UCMP 7418 (hypotype of 

Pseudotsuga glaucoides Axelr.) (p. 15). 

8. Stage 4+, Pinaceae gen. indet., specimen UCMP 7419B (hypotype of 

Pseudotsuga glaucoides Axelr.), x 2 (p. 15). 

9. Stage 4 +, Pinaceae gen. indet., specimen UCMP 7442 (paratype of 

Tsuga petranensis Axelr.), X 2 (p. 15). 

10. Pinus sp., atypical winged seed (p. 15). 

Atypical winged seed possibly belonging toP. crossii Knowlt., specimen 

UCMP 7299 (paratype of Picea coloradensis Axelr. ), x 2. 

11. Pinus aristata Engelm. 

Atypical winged seed, X 2. Introduced for comparison with preceding 

figure . 
US GEOLOGICAL SURVEY BULLETIN 1923 PLATE 2 

2 3 4 

\ 

., 

·}ii. 

7 

5 6 1 1 9 

10 

PINUS AND PINACEAE 
PLATE 3 

[All figures natural size unless otherwise indicated) 

FIGURES 1-3. Pinus remorata Mason. 

1. Ovulate cone preserved in longitudinal section. 

2. Exterior of ovulate cone presexved in three dimensions and with 10 attached cone scales. 

3. Same cone as in figure 2 but parts of cone scales removed, which represents abrasion stage 4 + . Potrero 

Canyon, California. Introduced for comparison. 

4. Pinus (Pinus) sp. 

Ovulate cone; basal two-thirds of cone represents abrasion stages 4 + and 5, and cleaned apical one-third of 

cone represents abrasion stage 1, showing typical apophyses, umbos, and mucros. Miocene, Turkey. 

Introduced for comparison. 

5. Pinus aristata Engelm. 

Winged seeds taken from a single cone. Introduced for comparison. 

Sa-f. Winged seeds atypical in development of seed and shape of wing. 

5g-5i. Winged seeds typical in development of seed and shape of wing. 

6. Picea sp. (p. 11 ). 

Specimen UCMP 7373, x2.5. 

7-13. Pinus crossii Knowlt. (p. 12). 

7. Winged seed, hypo type U CMP 7334, x 2.5. 

8, 9a. Staminate cone probably belonging toP. crossii, hypo type UCMP 9337b, x 2.5. 

9b. Winged seed, hypo type UCMP 9337a, x 2.5. 

10. Fascicle, hypotype UCMP 7221 (homeotype of P. florissantii Lesq.), x 2.5. 

11. Winged seed, hypotype UCMP 7220 (holotype of P. wasonii Axelr.), X 2.5. 

12. Fascicle, hypotype UCMP 7174 (hypotype of P. alvordensis Axelr.), x 2.5. 

13. Fascicle, hypotype UCMP 7361, x2.5. 
U.S. GEOLOGICAL SURVEY BULLETIN 1923 PLATE 3 

2 3 

4 

10 

12 13 

PINUS AND PICEA 
PLATE 4 

[All figures x 4 unless othetwise indicated) 

FIGURES 1-7, 9. Berberis coloradensis Axelr. (p. 15). 

1. Holotype UCMP 7527. 

2. Hypotype UCMP 7532 (paratype of B. riogrande Axelr.). 

3. Hypotype UCMP 7530 (holotype of B. riogrande Axelr.). 

4. Hypotype UCMP 7745 (paratype of Fend/era coloradensis Axelr.). 

5. Hypotype UCMP 7747 (paratype of F. coloradensis Axelr.). 

6. Hypotype UCMP 7665 (paratype of Philadelphus creedensis 

Axelr.). 

7. Hypotype UCMP 7656 (paratype of P. creedensis Axelr.). 

9. Hypotype UCMP 7849 (holotype of Vaccinium creedensis Axelr.). 

8, 10, 11. Mahonia aceroides (Knowlt.) Schorn & Wolfe (p. 17). 

Sa, lOa. Holotype USNM 36537 (holotype of Sterculia aceroides 

Knowlt.). 

8b, lOb. Hypotype USNM 36531. Fig. 8b, x 1. 

11. Hypotype UCMG 19749, x 1. 
U.S GEOLOGICAL SURVEY BULLETIN 1923 PLATE 4 

9 

10 

BERBERIS AND MAHON/A 
PLATES 

[All figure X 1.3 unless otheiWise indicated) 

FIGURES 1, 2, 5. Populus larsenii (Knowlt.) Schorn & Wolfe (p. 18). 

1. Hypotype UCMG 34060, x 1. 

2. Holotype USNM 36519 (holotype ofAlnus? larseni Knowlt.). 

5. Hypotypes UCMG 34061. 

3, 4. Jamesia capalanii Axelr. (p. 19). 

Hypotypes UCMG 34062, 340630. 

6, 7, 10, 11. Ribes lacustroides Axelr. (p. 20). 

6. Hypotype UCMG 27391 (holotype of R. creedensis Axelr.) 

7. Hypotype UCMG 34065. 

10. Hypotype UCMG 34066. 

8, 9. Ribes robinsonii Schorn & Wolfe (p. 22). 

8. Holotype UCMP 7817 (holotype of Physocarpus petiolaris Axelr.). 

9. Hypotype UCMG 34064. 
US GEOLOGICAL SURVEY BULLETIN 1923 PLATE 5 

POPULUS, JAMES/A, AND RIBES 
PLATE 6 

(All figures x 1.3] 

Figures 1-17. Ribes lacustroides Axelr. (p. 20). 

1. Hypotype UCMP 7581 (holotype of R. dissecta Axelr.). 

2. Holotype UCMP 7594. 

3. Paratype UCMP 7600. 

4. Paratype UCMP 7594. 

5. Hypotype UCMP 7626 (hypotype of R. stevenii Axelr.). 

6. Hypotype UCMP 7629 (homeotype of R. stevenii Axelr.). 

7. Hypotype UCMP 7603 (hypotype of R. stevenii Axelr.). 

8. Hypotype UCMP 7614 (hypotype of R. stevenii Axelr.). 

9. Hypotype UCMP 7602 (hypotype of R. stevenii Axelr.). 

10. Hypotype UCMP 7624 (homeotype of R. riogrande Axelr.). 

11. Paratype UCMP 7609. 

12. Hypotype UCMP 7837 (paratype of R. birdseyii Axelr.). 

13. Hypotype UCMP 7608 (homeotype of R. riogrande Axelr.). 

14. Hypotype UCMP 7625 (hypotype of R. riogrande Axelr.). 

15. Hypotype UCMP 7616 (homeotype of R. birdseyii Axelr.). 

16. Hypotype UCMP 7608 (homeotype of R. riogrande Axelr.). 

17. Hypotype UCMP 7632 (hypotype of R. webbii Wolfe). 
U.S. GEOLOGICAL SURVEY BULLETIN 1923 PLATE 6 

16 

RIBES 
PLATE 7 

(All figures X4 unless othexwise indicated) 

FIGURES 1-5. Eleiosina praeconcinna (Cocker.) Schorn & Wolfe (p. 23). 

1. Holotype UCMG 18536 (holotype of Solidago praeconcinna Cocker.), x 1. 

2. Hypotype UCMP 7389 (hypotype of Peraphyllum septentrionale [Lesq.] Axelr.). 

3. Hypotype UCMP 7505 (paratype of Salix creedensis Axelr.). 

4, 5. Hypotype UCMG 27387A, B (holotype of S. creedensis Axelr.). 

6. Eleiosina laevigata (L.) Rafin. Extant, Rock 23760, Yunnan Prov., China. Introduced for comparison. 

7. Peraphyllum ramosissimum Nutt. Extant, Gentry 143, Nevada, U.S.A. Introduced for comparison. 

8. Daphne septentrionale (Lesq.) MacG. 

Hypotype UCMP 3876, Florissant flora. Introduced for comparison. 
US GEOLOGICAL SURVEY BULLETIN 1923 PLATE 7 

3 

ELEIOSINA 
PLATE 8 

[All figures X 4 unless otheiWise indicated) 

FIGURES 1, 3, 5, 8, 9. Eleopoldia lipmanii (Axelr.) Schorn & Wolfe (p. 24). 

1, 3. Hypotype UCMG 34068. Fig. 3, x 1. 

5, 8. Hypotype UCMG 19786, Fig. 5, x 1. 

9. Hypotype UCMG 34069. 

2. Luetkea pectinata (Pursh) Kuntze. 

Extant, Thompson 5295, Washington, U.S.A. Introduced for comparison. 

4. Sanguisorba occidentalis Nutt. 

Extant, Fenis and Duthie 1254, Washington, U.S.A. Introduced for comparison. 

6. Geum ciliatum Pursh. 

Extant, Hitchcock 17993, Montana, U.S.A. Introduced for comparison. 

7. Fallugia paradoxa (D. Don) End!. 

Extant, Fosberg S3434, New Mexico, U.S.A. Introduced for comparison. 
U.S. GEOLOGICAL SURVEY BULLETIN 1923 PLATE 8 

ELEOPOLDIA 
PLATE 9 

(All figures X 4 unless othetwise indicated) 

FIGURES 1, 2. Ribes obovatum Schorn & Wolfe (p. 22 ). 

1. Holotype UCMP 7788 (homeotype of Holodiscus hameyensis Chan. & Axelr.). 

2. Paratype UCMG 34067, x 1.3. 

3, 4, 7, 8. Holodiscus stevenii Schorn & Wolfe (p. 25). 

3. Paratype UCMG 34071. 

4. Holotype UCMG 34070. 

7. Paratype UCMG 34072. 

8. Paratype UCMP7821 (hypo type of Rosa hilliae Lesq .). 

5. Sorbus potentilloides (Knowlt.) Axelr. (p. 27). 

Hypotype UCMG 19773. 

6. Crataegus creedensis Axelr. (p. 27). 

Holotype UCMP 7259B. 
U.S. GEOLOGICAL SURVEY BULLETIN 1923 PLATE 9 

5 

RIBES, HOLODISCUS, SORB US, AND CRA TAEGUS 
PLATE 10 

(All figures x 4 unless otherwise indicated] 

FIGURES 1, 2, 8, 9. Crataegus holmesii Lesq. [ = Cercocarpus holmesii (Lesq.) Axelr.] (p. 29). 

Silver Cliff, Colorado. Introduced for comparison. 

1. Syntype USNM 313296. 

2. Syntype USNM 313294. 

8. Syntype USNM 313295. 

9. Lectotype USNM 313299. 

3-6, 15, 16. Cercocarpus henricksonii Schorn & Wolfe (p. 28). 

3. Holotype UCMG 34073, x 1.3. 

4. Paratype UCMG 34074, x 1.3. 

5. Paratype UCMG 34075, x 1.3. 

6. Paratype UCMG 34076. 

15, 16. Paratype UCMG 34077. Fig. 15, x 1.3. 

7. Potentilla creedensis (Axelr.) Schorn & Wolfe (p. 27). 

Hypotype USNM 422590, x 1.3. 

10, 11. Andromeda linearifolia Lesq. [ = Cercocarpus linearifolius (Lesq.) Axelr.] (p. 30). 

Silver Cliff, Colorado. Introduced for comparison. 

10. Lectotype USNM 313291. 

11. Syntype USNM 23528. 

12-14. Cercocarpus nanophyllus Schorn & Wolfe (p. 29). 

12. Holotype UCMG 34078. 

13. Paratype UCMG 34079. 

14. Paratype UCMG 34080. 
U.S. GEOLOGICAL SURVEY BULLETIN 1923 PLATE 10 

3 

9 

CERCOCARPUS AND POTENTILLA 
PLATE 11 

(All figures X 1.3 unless otheiWise stated] 

FIGURE 1. Potentilla creedensis (Axelr.) Schorn & Wolfe (p. 27). 

Holotype UCMG 19757 (holotype of Ranunculus creedensis Axelr.). 

2, 5-7. Prunus creedensis Axelr. (p. 31). 

2. Hypotype UCMG 19774 (holotype of Sambucus longifolius Axelr.), x 4. 

5. Hypotype UCMG 34082. 

6. Hypotype UCMG 34083. 

7. Hypotype UCMG 34084. 

3, 8. Rosoideae, genus and species indetermined (p. 30). 

3. Specimen UCMG 19766. 

8. Specimen UCMG 34081. 

4, 9. Osmaronia? stewartiae (Axelr.) Schorn & Wolfe (p. 30). 

Holotype UCMG 19778 (holotype of Arbutus stewartii Axelr.). Fig. 9, X 4. 
US GEOLOGICAL SURVEY BULLETIN 1923 PLATE 11 

4 

POTENTILLA, PRUNUS, ROSOIDEAE, AND OSMARONIA? 
PLATE 12 

(All figures natural size unless othetwise indicated) 

FIGURES 1, 2, 7. Catalpa coloradensis (Axelr.) Schorn & Wolfe (p. 33). 

1. Paratype UCMP 7521. 

2. Hypotype UCMG 34085, x 0.6. 

7. Paratype UCMP 7522. 

3. "Chilopsis coloradensis Axelr." (p. 35). 

Name of dubious value. Holotype UCMP 7852, x 4. 

4, 6. "Quercus creedensis Axelr." (p. 34). 

Name of dubious value. 

4. Holotype UCMP 7516, x 4. 

6. Paratype UCMP 7515, x 4. 

5. Prunus creedensis Axelr. (p. 31) 

Hypotype USNM 422593, x 1.3. 

8. "Fraxinus creedensis Axelr." (p. 35) 

Name of dubious value. Holotype UCMG 19777, x 4. 
U.S. GEOLOGICAL SURVEY BULLETIN 1923 PLATE 12 

3 

8 

CATALPA AND PRUNUS 
PLATE 13 

[All figures natural size unless otherwise indicated) 

FIGURES 1, 2. Group of achenes of uncertain affinity (p. 28). 

Specimens UCMP 7781A, B (paratype of Fallugia lipmanii Axelr.), x 4. 

3. Eleopoldia lipmanii (Axelr.) Schorn & Wolfe (p. 24). 

Hypotype UCMP 7445 (holotype of Juniperns gracillensis Axelr.), x 5. 

4. Pinus crossii Knowlt. (p. 12). 

Hypotype UCMG 34154. 
US. GEOLOGICAL SURVEY BULLETIN 1923 PLATE 13 

PINUS, ELEOPOLDIA , AND ACHENES 

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