331

Int. J. Plant Sci. 161(2):331344. 2000.

2000 by The University of Chicago. All rights reserved.
1058-5893/2000/16102-0016$03.00
CONES, SEEDS, AND FOLIAGE OF TETRACLINIS SALICORNIOIDES (CUPRESSACEAE) FROM
THE OLIGOCENE AND MIOCENE OF WESTERN NORTH AMERICA: A GEOGRAPHIC
EXTENSION OF THE EUROPEAN TERTIARY SPECIES
Zlatko Kvacek, Steven R. Manchester,
1
and Howard E. Schorn
Charles University, Faculty of Science, Albertov 6, 12843 Prague 2, Czech Republic; Florida Museum of Natural History, University of Florida,
Gainesville, Florida 32611, U.S.A.; and Museum of Paleontology, University of California, Berkeley, California 94720, U.S.A.
The cupressaceous genus Tetraclinis is recognized fromthe Oligocene and Miocene of western North America
on the basis of co-occurring seed cones, seeds, and foliage branches. Morphological and anatomical compar-
isons with the two previously recognized European Tertiary species indicate that the North American specimens
are morphologically inseparable from Tetraclinis salicornioides (Unger) Kvacek. The North American taxon
is treated as a new variety, T. salicornioides (Unger) Kvacek var. praedecurrens (Knowlton) comb. et stat. nov.,
and is distinguished from the European representatives, T. salicornioides (Unger) Kvacek var. salicornioides,
by slight anatomical differences in the leaf epidermis. Although cones and seeds of the fossil species are closely
similar to those of extant Tetraclinis articulata, the foliage is more spreading, composed of attened segments
with fused facial and lateral leaves that are apparently adaptive for a more mesic climate. The recognition of
T. salicornioides in western North America along with the absence of Tetraclinis in the fossil and recent ora
of eastern Asia provide evidence for communication of the species across the North Atlantic during the early
or middle Tertiary.
Keywords: cones, Cupressaceae, Europe, foliage, North America, paleobotany, Tertiary.
Introduction
The cupressaceous genus Tetraclinis Masters has a single
extant species, Tetraclinis articulata (Vahl) Masters, that is
native to warm, summer-dry climates of northern Africa,
Malta, and southern Spain. In addition, the genus has an ex-
cellent fossil record in the Tertiary of Europe based upon fossil
cones, seeds, and foliage (Kvacek 1989; Kovar-Eder and Kva-
cek 1995; Mai 1996). Two fossil species have been recognized:
Tetraclinis brachyodon (Brongniart) Mai et Walther (Early Eo-
cene to Early Pliocene) and Tetraclinis salicornioides (Unger)
Kvacek (Middle Eocene to Early Pliocene), although some au-
thors merge these species into a single entity (Mai 1996, 1997).
In this article we recognize another representative of Tetraclinis
from the Oligocene and Miocene of North America on the
basis of associated cones, seeds, and foliage similar to the Eu-
ropean fossil T. salicornioides. This occurrence, as recently
recorded by Meyer and Manchester (1997), conrms that Te-
traclinis was formerly distributed outside of Europe and north-
ern Africa.
Tetraclinis has bilaterally symmetrical seed cones composed
of four slightly uneven valvate scales and seeds that are basally
cordate with a pair of large membranous wings. Leaves of the
single extant species are arranged in pseudowhorls and are
very reduced, rather like those of incense cedar (Calocedrus
decurrens [Torr.] Florin). However, in the Tertiary of Europe,
two kinds of foliage have been distinguished. Foliar branchlets
1
Author for correspondence; e-mail steven@mnh.u.edu.
Manuscript received March 1999; revised manuscript received October 1999.
of T. brachyodon (Brongniart) Mai et Walther have reduced
leaves that are difcult to distinguish from those of the extant
species. Those of T. salicornioides are broader and more at-
tened and appear to be adapted to more mesic conditions. The
generic identity of these foliage types has been conrmed by
the recovery of specimens that showa physical connection with
the characteristic seed cones of Tetraclinis.
In this article we review the taxonomy and distribution of
the North American population; provide a full description of
the cones, seeds, and foliage (including newdata fromcuticular
analyses); and compare the North American population with
other fossil Cupressaceae, including European representatives
of Tetraclinis. Various morphological and molecular analyses
corroborate the merger of the Cupressaceae and Taxodiaceae
into one group (Eckenwalder 1976; Brunsfeld et al. 1994; Ste-
fanovic et al. 1998). To avoid nomenclatural misunderstand-
ing, we use the term Cupressoids in the sense of Cupres-
saceae s.s. in the following text.
Material and Methods
Fossil leaves, seed cones, and seeds were observed in col-
lections from the U.S. National Museum (USNM) in Wash-
ington, D.C.; the University of California Museum of Pale-
ontology (UCMP) in Berkeley; and the Florida Museum of
Natural History (UF) in Gainesville. Oligocene specimens are
from the Lost Creek, Lyons, and Willamette oras of Oregon
(Meyer and Manchester 1997) and the Gumboot Mountain
ora of Washington (UCMP loc. PA 399). Miocene specimens
are from the Eagle Creek (Chaney 1920), Mollala (Chaney
and Axelrod 1959), and Collawash (U.S. Geological Survey
332 INTERNATIONAL JOURNAL OF PLANT SCIENCES
Table 1
Localities for Tetraclinis salicornioides var. praedecurrens Showing
Co-occurrences of Foliage, Cones, and Seeds
Locality Foliage Cones Seeds
Oligocene:
Lost Creek I, Oregon . . . . . . . . . . . . . . . . . . . x x x
Lost Creek II, Oregon . . . . . . . . . . . . . . . . . . . x x x
Willamette, Oregon . . . . . . . . . . . . . . . . . . . . . x ) x
Lyons, Oregon . . . . . . . . . . . . . . . . . . . . . . . . . . . x ) x
Gumboot Mountain, Washington . . . . . . x ) )
Miocene:
Potlatch Creek, Idaho . . . . . . . . . . . . . . . . . . . x ) x
Oviatt Creek, Idaho . . . . . . . . . . . . . . . . . . . . . x ) )
Molalla, Oregon . . . . . . . . . . . . . . . . . . . . . . . . . x ) )
Collawash, Oregon . . . . . . . . . . . . . . . . . . . . . . x ) )
Eagle Creek, Oregon . . . . . . . . . . . . . . . . . . . . x ) )
Spokane, Washington . . . . . . . . . . . . . . . . . . . x ) )
Grand Coulee, Washington . . . . . . . . . . . . . x ) )
[USGS] loc. 9526; Peck et al. 1964) localities of western
Oregon, the Potlatch Creek (Brown 1935) and Oviatt Creek
(Boyd 1985) oras of Idaho, and the Latah (Knowlton 1926)
and Grand Coulee (Berry 1931) oras of Washington. A list
of the localities and the organs that have been collected is
provided in table 1.
Comparative material of Tetraclinis salicornioides var. sal-
icornioides and T. brachyodon from the Tertiary of Europe
was examined in collections the Museum National dHistoire
Naturelle in Paris (MNHN; collections of Brongniart and Sa-
porta), the Museumfu r Naturkunde der Humboldt-Universita t
in Berlin (collection of D. H. Mai), the Museum Joanneum in
Graz, Austria (collection of Unger), the Universita t Graz (col-
lection of Ettingshausen) fromthe National Museumin Prague
(NM), and Charles University (Faculty of Science) in Prague
(PRC). These materials were supplemented by material from
the other institutions acknowledged in Bu zek et al. (1976),
Knobloch and Kvacek (1976), and Kvacek (1989).
Cuticles were prepared for light microscopy using specimens
from the Oligocene Willamette ora and the Miocene Latah,
Potlatch Creek, and Mollala oras of Idaho. The cuticle was
macerated with routine methods (Knobloch and Kvacek 1976),
using Schultzes solution and KOH (5%), and the cuticle was
then studied with transmitted light and scanning electron mi-
croscopy (SEM). Isolated cuticles for SEM were prepared by
spreading the fragments on the moistened emulsion surface of
4-mm-square pieces of photographic lm that were subse-
quently glued to the aluminum stubs.
Systematics
FamilyCupressaceae
GenusTetraclinis Masters
Tetraclinis salicornioides (Unger) Kvac ek var.
praedecurrens (Knowlton) comb. et stat. nov.
(Fig. 1A1E, 1G, 1H; Fig. 2A2G, 2I2L;
Fig. 3A3E; Fig. 4A, 4C, 4D)
Basionym. Libocedrus praedecurrens Knowlton 1926,
USGS Prof. Pap. 140:28, pl. 8, g. 8 [foliage].
Synonyms. Fokienia praedecurrens (Knowlton) Chaney
and Axelrod 1959, Carnegie Inst. Wash. Publ. 617:145, pl.
14, g. 8 [foliage].
Callitris potlatchensis Brown 1935, J. Paleontol. 9:575, pl.
67, g. 16. Brown 1940, J. Wash. Acad. Sci. 30:347, g. 6
[seed].
Tetraclinis potlatchensis (Brown) Meyer and Manchester
1997 (potlachensis), Univ. Calif. Publ. Geol. Sci. 141:64, pl.
3, gs. 19 [seed and cone].
Fokieniopsis praedecurrens (Knowlton) Meyer and Man-
chester 1997, Univ. Calif. Publ. Geol. Sci. 141:64, pl. 3, gs.
1014 [foliage].
Description. Cone wide-ovate, valvate; cone scales ap-
parently four (but sometimes only three preserved) in opposing
pairs, incompletely preserved, dimorphic, one pair ca. 1.3
times wider than the other, thick and woody, widely ovate,
length 78 mm, width 912 mm, base cordate, apex rounded;
abaxial surface of cone scale with a mucro positioned ca. one-
third of the distance from the base to the apex of scale; surface
of scale wrinkled, except for a smooth, broadly elliptical area
(fused bract) at the base surrounding the mucro, hemispherical
in outline; adaxial surface of scales with ne striations radi-
ating from the base.
Seed with two wings, symmetrical, reniform to broadly
ovate, base cordate to sometimes truncate, apex obtuse to
rounded, overall width 712 mm, length 47.5 mm; wings
equal, much wider than seed body, thin, lateral margins
rounded; seed body ovate, apex acute, base rounded, length
37 mm, width 23.5 mm, longitudinal striae radiating from
basal attachment scar and converging toward bilobed micro-
pylar tip and demarcating elliptical resin vesicles.
Foliage twigs with mostly opposite branching. Branchlets
attened with four-ranked dimorphic scale leaves borne in
pseudowhorls. Leaves dimorphic, facial and lateral scalelike
leaves with rounded to bluntly mucronate apices. Facial and
lateral leaves of each pseudowhorl fused (usually) most of their
length to form a cladode-like dorsiventrally attened segment.
Lateral leaves do not overlap between adjacent nodes, and the
facial leaves overlap only slightly or not at all. Segments cov-
ered by stomata on both surfaces, cuticular membrane thick,
leaves amphistomatic. Abaxial side of fused leaves composed
of several longitudinal, vaguely differentiated nonstomatal and
stomatal areas. Ordinary cells of nonstomatal areas quadran-
gular-polygonal, usually much elongated, 35100 mmlong and
2030 mm wide. Anticlinal walls very regularly nely undulate
(to straight); sinuses of the undulation with papilloid thick-
enings. Longitudinal stomatal areas containing short rows of
closely spaced stomata or irregularly oriented stomatal groups
separated by nonstomatal areas. Stomata monocyclic, irreg-
ularly oriented, stomatal pit quadrangular to polygonal, 2030
mm wide, including a thick Florin ring. Ordinary cells between
stomata triangular to polygonal, more or less isodiametric,
(12) 3037 mm across or quadrangular, elongate, 1530 mm
wide and up to 70 mm long. More or less distinct papillae in
stomatal areas, broad prominent papillae near the segment
base. Small quadrangular crystal cavities in the cuticular
membrane observed near the terminal part of a foliar segment.
Free tips or free margins of the scale leaves with marginal
papillae forming prominent crests. Cuticles of adaxial side of
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EK ET AL.CUPRESSACEAE IN THE NORTH AMERICAN TERTIARY 333

Fig. 1 Cones and seeds of Tetraclinis. AE, Tetraclinis salicornioides var. praedecurrens from the Oligocene of Lost Creek Reservoir, Oregon,
#2.5. A, Cone in lateral view showing apically rounded basally cordate cone scale and the attachment to stalk with free-tipped foliar segments;
USNM 450636. B, The same cone as in g. 1A, with portion of the facing cone scale removed to show the pair of lateral cone scales behind.
C, An opened cone transversely compressed, with three of the cone scale visible; UF 10152. D, One large lateral cone scale (right) and a pair
of opened cone scales (top and bottom), showing position of the mucro; UF 10151. E, Counterimpression of the specimen in g. 1D, showing
thick, woody nature of the cone scales. F, Extant Tetraclinis articulata, apical view, showing the four cone scales, Algeria; US 2039727, #3.
G, Seed showing two wings attached to ovate seed body, proximal ends of the wings truncated by folding; UF 10163, #3. H, Another seed
showing longitudinal resin vesicles on the seed body and large lateral wings; UF 10164, #3. I, European specimen for comparison: cones of T.
salicornioides var. salicornioides from Suletice, Czech Republic; NM G7535A, #2.5. J, Cone of extant T. articulata (Vahl) Masters from
Marrakech, Morocco (PRC), lateral view, #2.5. K, Seed of T. salicornioides var. salicornioides from Markvartice, Czech Republic; NM G2785,
#4. L, M, Seeds from the cone in g. 1J, #2.5.
334 INTERNATIONAL JOURNAL OF PLANT SCIENCES
Fig. 2 Foliar branches of T. salicornioides var. praedecurrens and salicornioides, all #2.5. AG, T. salicornioides var. praedecurrens. A, Type
of Knowlton 1926, pl. 8, g.8, p. 28, from Miocene of Latah ora, Spokane, Washington; USNM 36873. B, Branch from the Oligocene Lyons
ora, Oregon, showing opposite ultimate branches; USNM 450640. CG, From Oligocene of Lost Creek Reservoir, Oregon. C, Twig showing
opposite branching with cuneiform medial segments giving rise to ultimate branches with simple, obovate segments; UF 10158. D, Elongate
segments from proximal part of shoot showing arrested lateral growth and free tips of the lateral leaves; UF 10161. E, Series of elongate foliar
segments from the proximal part of a twig; UF 10160. F, Similar twig with more robust segments; UF 10159. G, Succession of three medial
cuneiform foliar segments; UF 10153. H, T. salicornioides var. salicornioides shoot from Kundratice showing three kinds of foliar segments:
narrow elongate proximal segments and cuneiform medial segments giving rise to simple lateral ultimate segments; NM G 2172. IL, Isolated
foliar segments of T. salicornioides var. praedecurrens from Lost Creek Reservoir, Oregon. IK, Cuneiform segments of the branch junction,
showing ve main veins. I, UF 10154. J, UF 10155. K, UF 10157. L, Segment of an ultimate branch with three veins; USNM 450637.
KVAC

EK ET AL.CUPRESSACEAE IN THE NORTH AMERICAN TERTIARY 335

Fig. 3 Cuticle of Tetraclinis salicornioides. A, Medial cladode-like segment of T. salicornioides var. praedecurrens from which cuticle was
removed for epidermal anatomy, from Potlatch Creek, Idaho; UF10887, #5. B, A lateral segment, cleared showing incomplete fusion of leaves,
from Oviatt Creek, Idaho; UF 18580-31378, #5. C, Free tip of the lateral leaf, lying on dorsal surface, showing adaxial and abaxial cuticle
folded together with stomata on each; USNM 36873 (prep. 1), #40. D, Facial leaf, abaxial surface showing groups of closely spaced stomata;
USNM 36873 (prep. 1), #200. E, Cuticle from near the base of segment in g. 3A, showing longitudinally aligned epidermal cells and the
arrangement of stomata in rows, #200. F, T. salicornioides var. salicornioides from the Oligocene site Markvartice, western Czech Republic,
for comparison with the North American specimens; PRC MR 134-B, #200. G, Same as g. 3E, by phase-contrast light microscopy, showing
the papillate nature of the epidermal cells, #400.
leaves thin, nely papillate, densely covered with stomata of
the same type as that seen on the abaxial surface.
Holotype. USNM 36873, railroad cut in Spokane, Wash-
ington, Miocene Latah ora, Washington (Knowlton 1926),
g. 2A; g. 3C, 3D.
Additional specimens. OligoceneUF 10153-10158,
10161, 10575; USNM 450637; UF 10162-10165; USNM
42333; UF 10151, 10152, 10576; USNM450636 (Lost Creek,
Oreg.; UF localities 243, 244); UF 9086; USNM 450641,
450644 (Willamette ora, Oreg.; UF locality 15784). Mio-
ceneUF 10887 (Potlatch Creek, Julietta, Idaho; UF locality
18597); UF 26762, 26763 (Lower Salmon River, Idaho; UF
locality 18598); UF 31378 (Oviatt Creek, Idaho; UF locality
18580); UCMP 4421 (Mollala, Oreg.); Field MuseumChicago
22335 (Eagle Creek, Oreg.).
Additional localities. In addition to the localities indicated
above, specimens are known from the Miocene oras of Col-
lawash, Oregon, and Grand Coulee, Washington (Chaney and
Axelrod 1959).
Discussion. In the above description, we combine the
seeds previously assigned to C. potlatchensis Brown with the
foliage previously called F. praedecurrens (Knowlton) Chaney
and Axelrod, together with the associated cones, under the
new combination T. salicornioides var. praedecurrens. The
combination of these organs under the same name is justied
by the co-occurrence of the seeds and foliage at ve localities,
by the co-occurrence of the cones, seeds, and foliage at two
localities (table 1), and by the previously recognized co-
occurrence and physical attachment of similar foliage, cones,
and seeds of T. salicornioides in the Tertiary of Europe (Kvacek
1989).
Brown (1935, 1940) was the rst to recognize the cupressoid
afnity of the seeds from the Miocene of Idaho and the Oli-
gocene of Oregon, and he named them C. potlatchensis. His
generic assignment was based upon a correct comparison with
the extant Tetraclinis articulata (Vahl.) Masters, which was
previously placed under the synonym Callitris quadrivalvis
Ventenant. Although not found in situ within the cones, the
associated seeds coincide in size and conguration with the
area for seed attachment within the fossil cones, thereby sup-
porting our conclusion that the cones and seeds represent the
same species. The cones and seeds are remarkably similar to
336 INTERNATIONAL JOURNAL OF PLANT SCIENCES
Fig. 4 Cuticle of Tetraclinis salicornioides in SEM. A, T. salicornioides var. praedecurrens. Inner side of cuticle showing anticlinal walls of
ordinary cells with thickenings, Potlatch Creek, Idaho; UF 10887, #500. B, T. salicornioides var. salicornioides from the Early Miocene of
Plesna, western Czech Republic, for comparison with the North American specimen, inner side of cuticle showing strongly papilla-like thickenings
on anticlinal walls of ordinary cells and a stoma; PRC V-146-112, #500. C, T. salicornioides var. praedecurrens, outer side of the cuticle,
showing stomata and papillae; UF10887, #500. D, Same, enlarged to show stomatal aperture and surrounding Florin ring; UF
10887, #1500.
those of T. salicornioides (Unger) Endlicher from the Tertiary
of Europe (Kvacek 1989; Kvacek and Walther 1995).
Only four seed cone specimens have been recovered from
the North American record, and they are all from the Oli-
gocene Lost Creek locality of the Bridge Creek ora in Oregon
(Meyer and Manchester 1997). One is compressed laterally
(g. 1A, 1B), and the others are compressed transversely or
slightly obliquely with the scales opened (g. 1C1E). One of
the specimens shows only three enlarged cone scales, indicating
that one was arrested in development, as is also sometimes
observed in the European fossil and extant cones of Tetraclinis
(Zablocki 1928, pp. 188189).
Cones with four dimorphic valvate cone scales provide de-
cisive support for the assignment of these fossils to Tetraclinis.
The fossil seeds also compare favorably with those of extant
Tetraclinis in their bisymmetry, cordate base, large wings, and
resin vesicles on the seed body. As in the European fossils of
T. salicornioides (Kvacek 1989), the North American seeds are
less prominently cordate than are the seeds of extant T. arti-
culata. The lower position of the mucro on the abaxial surface
of the cone scale provides an additional character that distin-
guishes these fossils from the modern species and that under-
scores the similarity with the European fossils. In extant T.
articulata, the mucro usually occurs in the upper one-fourth
of the cone scale, near the apex. In the two cones in which
this can be observed among the Lost Creek specimens (g. 1A,
1D), the mucro is situated ca. one-third of the distance from
the base toward the apex of the cone scale. In the European
representatives of T. salicornioides, the mucro ranges in po-
sition from the lower one-third (Kelber and Gregor 1987, pl.
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EK ET AL.CUPRESSACEAE IN THE NORTH AMERICAN TERTIARY 337

3, g. 3) to a maximum of one-half of the length of the cone
scale (Schloemer-Ja ger 1960, pl. 1, g. 15).
Foliage of T. salicornioides var. praedecurrens was formerly
assigned to Fokienia based on similarities in leaf morphology
by Chaney and Axelrod (1959). However, in the living Fok-
ienia hodginsii Dunn, the branch segments are concave on the
underside (rather than biconvex), the leaves are not fused, and
the apices of facial and lateral leaves are more acute (McIver
and Basinger 1990, pp. 16131614, gs. 16, 17). As observed
by Meyer and Manchester (1997), the fossil foliage from these
North American localities is morphologically indistinguishable
from that of the widespread European Tertiary species T.
salicornioides.
Until being recognized as Tetraclinisa recognition that was
inuenced by the attached cones and associated seeds (Kvacek
1989)the European foliage specimens were considered to
belong to an extinct genus, for which the names Hellia Unger
[nom. invalid: generic diagnosis lacking; three species were
attributed to the genus, but a generitype was not designated]
(e.g., Mai 1963; Friis 1977) and Libocedrites Endlicher (e.g.,
Knobloch and Kvacek 1976; Mai and Walther 1978; Pala-
marev et al. 1991; Wilde and Frankenha user 1998) had long
been applied. We use the term cladode-like in reference to the
foliar branch segments. They are distinguished from true clad-
odes (derived frombranches without leaves) by their derivation
from leaves plus the branch. The cladode-like segments consist
of mutually fused dimorphic scale leaves borne in pseudo-
whorls. The nearly complete fusion of lateral and facial leaves
does not occur among extant Cupressaceae (not even in the
extant species of Tetraclinis) but does occur in both the Eu-
ropean and North American populations of T. salicornioides.
The cladode-like branches typically disaggregate at succes-
sive nodes into short segments that may be found dispersed in
the sediment (g. 2I2L; g. 3A, 3B). These internode seg-
ments occur in four forms: the rst are simple segments from
the ultimate portions of twigs and branches (e.g., g. 2A2C).
They are elliptical or obovate and have three main veins (e.g.,
g. 3B). The second kind are medial segments that represent
the junctions at which one or two side branches arise (e.g.,
g. 2A, 2H2K; g. 3A). They are obovate to wedge shaped,
with a jagged distal end and ve visible veins, the outer pair
of which are similar to those of the ultimate segments and the
inner pair of which leads to the branches. The third kind are
narrow segments situated in the proximal portion of the shoot.
They are approximately parallel sided over most of their length
but are abruptly enlarged distally, with free tips on the lateral
leaves, which subtend arrested axillary branches (g. 2D2F).
The fourth kind of segment, which is typically situated below
the cones, is shorter, less dimorphic, and has free tips (g. 1A,
1B). As a result of fusion, the boundaries between individual
leaves cannot be discerned further down the segment, except
in rare cases of incomplete fusion (e.g., g. 3B; Friis 1977)
and in branches bearing cones (Kvacek 1989).
Friis (1977) made detailed comparisons of T. salicornioides
(Unger) Kvacek (as Hellia salicornioides) foliage segments with
extant cupressoid genera and observed some agreement with
extant T. articulata, both in terms of epidermal structure and
in the external and internal morphology. She noted, however,
that the extant species is distinguished by markedly xero-
morphic characters with deeply depressed stomatal bands and
without stomata on exposed surfaces. Tetraclinis salicornioides
has wider leaves covered by stomata on the abaxial sides, and
it bears dense stomata on the adaxial sides of the free parts
of leaves. These features are consistent with the mesic habitat
that can be inferred for both varieties of T. salicornioides,
based upon the oristic associations in which they occur.
We consider it appropriate to place the North American
material in the same species as the European fossils, T. sali-
cornioides, because the morphology of the foliage and of the
seeds fully coincides. Although the geographic disjunction
might lead one to expect distinct species, the morphological
differences are slight. The cones tend to be slightly larger in
the North American specimens, but this may be attributed to
differences in preservation type (impression vs. compression),
and there is overlap in the dimensions (cone width 912 mm
vs. 310 mm). Without the aid of leaf cuticular characters
(mainly stomatal arrangement), it would be difcult to make
any distinction between the North American and European
populations.
Varietal status for the North American material is supported
by differences in epidermal anatomy. In the European material,
the arrangement and shape of epidermal cells are less regular
than in the North American Miocene material. The anticlinal
walls usually bear thickenings and are more intensively un-
dulate (gs. 3F, 4B vs. g. 3D, 3E, and g. 4A), and the
periclinal walls more often have distinct papillae. Stomata are
generally more widely scattered. However, the epidermal char-
acters of the Oligocene foliage (from the Willamette ora) are
less different from the type European representatives of the
species. This indicates an increasing divergence of the Euro-
pean and American populations that may have arisen follow-
ing their geographical separation.
Conspectus of the Fossil Representatives of Tetraclinis
Some authors (Mai 1997, 1998; Ferguson et al. 1998) main-
tain that only a single species, Tetraclinis brachyodon (includ-
ing Tetraclinis salicornioides), existed in the European Tertiary.
However, in our investigation of numerous fossil occurrences
of Tetraclinis, we conrmed that there are indeed distinguish-
ing criteria (in both the seed cones and the foliage) that support
recognition of the two species, as indicated in the key and
descriptions below.
Diagnostic key to fossil Tetraclinis species:
I. Branching mostly alternate, leaf fusion incomplete, sto-
mata lacking on abaxial side, bract of the seed cone elongate,
mucro on abaxial surface of seed cone scale typically subter-
minal ) Tetraclinis brachyodon (Brongniart) Mai et Walther
II. Branching mostly opposite, leaf fusion mostly complete,
stomata present on abaxial side, bract of the seed cone broader
than long, mucro typically subcentral to subbasal )Tetraclinis
salicornioides (Unger) Kvacek
Tetraclinis brachyodon (Brongniart) Mai et Walther (syn.
Thuytes callitrina Unger, Callitrites brongniartii
Endlicher, nom. illegit. super.)
Morphology and anatomy. This species is based on sterile
twigs from Middle Eocene strata in ParisMont Rouge/Cha-
tillon (lectotype: Brongniart 1822, pl. 5, g. 3A, no. 78, Mus.
338 INTERNATIONAL JOURNAL OF PLANT SCIENCES
Nat. Paris; see g. 5E). Its leaf segments are nearly isomorphic,
very slender, arranged in pseudowhorls with 2 (4) subparallel
longitudinal grooves marking the junctions of incompletely
fused scale leaves (g. 5E, 5F, 5K, 5L). Branching is alternate
and rarely opposite. The abaxial epidermis is composed of
nonpapillate straight-walled cells, with stomata conned
mostly to the grooves between the scale leaves (Givulescu
1975; Kvacek 1989; our g. 5M). Seed cones (from various
localities, including Armissan, Aix-en-Provence, St. Zacharie,
Radoboj, and Ha ring; e.g., our g. 5A5D, 5G) are broadly
truncate conical to globular, attached singly to the twigs; the
fused bract elongate, ascending often more than two-thirds of
the cone scale length and mostly indistinct, with a subterminal
to rarely subcentral umbo. Seeds (g. 5H, 5I) bilaterally
winged, deeply to shallowly cordate, rarely truncate.
Occurrence and stratigraphic range. Europe to Trans-
caucasia, Early Eocene to Early Pliocene.
Tetraclinis salicornioides (Unger) Kvac ek (syn. Libo-
cedrites salicornioides [Unger] Endlicher)
Morphology and anatomy. This species is typied by ster-
ile twigs from the Miocene of Radoboj (lectotype: Unger 1841,
pl. 2, g. 1, no. 76680, Landesmuseum Joanneum, Graz, Aus-
tria). Leaf segments are more or less broad and well attened
and remarkably polymorphic according to their position
within sprays. They consist of fully fused scale leaves in pseu-
dowhorls, with grooves between lateral and facial leaves barely
visible in the uppermost part of the segments (g. 2B2L) or,
more rarely, reaching toward the segment base (e.g., Friis 1977;
our g. 2A). Branching is opposite to rarely alternate. Abaxial
epidermis composed of more or less strongly papillate cells
with nely undulate anticlinal walls and stomata in irregular
groups or longitudinal lines over the whole surface of the seg-
ments (gs. 3B3G, 4A4D). Seed cones are rarely found in
attachment to the twigs (from the localities of Haselbach,
Suletice-Berand, Flo rsheim, and Kreuzau), singly or in pairs.
They are broadly ovate (broader than long), and the fused
bract is broader than long and roundish to broadly cordate
with a subbasal to subcentral umbo (g. 1A1F). Bilaterally
winged seeds are shallowly cordate to truncate (g. 1G, 1H,
1K).
Occurrence and stratigraphic range. Variety salicornioides
is distributed from Europe to Transcaucasia, Middle Eocene
to Early Pliocene (Kvacek 1989). The second variety, prae-
decurrens, discussed below, is distributed in western North
America.
Tetraclinis salicornioides (Unger) Kvac ek var.
praedecurrens (Knowlton) Kvac ek and
Manchester, comb. et stat. nov. (syn.
Libocedrus praedecurrens Knowlton,
Callitris potlatchensis Brown)
Morphology and anatomy. This variety is based on a ster-
ile twig fromthe Miocene Latah ora in Washington (holotype:
Knowlton 1926, pl. 8, g. 8, USNM 36873; our g. 2A; g.
3C, 3D). It differs from the type variety only by its epidermal
anatomy, consisting of more regular arrangement and shape
of epidermal cells, less distinctly thickened and less intensively
undulate anticlinal walls, and stomata that are more closely
spaced and in more regularized rows (gs. 3E, 4A).
Occurrence and stratigraphic range. Western United
States, Oligocene and Miocene (Meyer and Manchester 1997).
Contrary to the opinions of some authors (Friis 1977;
Palamarev et al. 1991; Wilde and Frankenha user 1998; E. M.
Friis, personal commmunication, 1998), the above differences
probably do not warrant the assignment of T. salicornioides
to an extinct genus (i.e., Libocedrites Endlicher). The combi-
nation of quadrivalved seed cones with noncolumellate di-
morphic cone scales and with bilaterally broadly winged seeds
is unique among the Cupressoids and is diagnostic of extant
Tetraclinis (Kra usel 1938; Mai and Walther 1978; Kvacek
1989; Martinetto 1995; Mai 1997; Meyer and Manchester
1997). We chose to consider the unusual cladode-like foliage
and branching pattern of T. salicornioides as features of sub-
ordinate taxonomic value. In fact, some rare foliage forms
mentioned above (g. 2A) intergrade morphologically with the
typical foliage of T. brachyodon. The latter fossil species
matches extant T. articulata in most respects, both in its foliage
and seed cones. The close linkage between T. salicornioides,
T. brachyodon, and T. articulata weakens the argument for
treating any part of this complex as an extinct genus.
Cones of Cupressinites curtus Bowerbank (Tetraclinis curta
[Bowerbank] Mai 1997) from the Early Eocene London Clay
ora appear, based on the published descriptions and illustra-
tions (Bowerbank 1840), to be similar to Tetraclinis, and with
additional study, they may provide important information
about the early differentiation of the Tetraclinis cone mor-
phology. D. H. Mai (personal communication, 1997) studied
the available material and did not hesitate to include it into
Tetraclinis. However, these cones are reported to be up to ve-
scaled (Collinson 1983), and no information on their foliage
is available.
Cones and Seeds of Cupressaceae
The small number of cone scales and their decussate (rather
than helical) arrangement, combined with their highly fused
bract and scale, are features that conrm the afnities of these
fossils with the Cupressoids. Seed cones of extant Cupressoids
range from more or less globular to elongate and may have
peltate, imbricate, or valvate cone scales that may be woody
to leathery or eshy. Verticillate arrangement of scales occurs
in some Callitroidae of the Southern Hemisphere. The com-
bination of globose cones and four valvate, unequal, woody
cone scales, combined with the peculiar broadly elliptic and
basally cordate seeds, indicates an afnity to extant Tetraclinis.
Seeds of Cupressaceae are relatively consistent within each
natural genus and provide characters of diagnostic value, par-
ticularly in terms of the disposition and form of the wings.
The most common type of seed has two narrow wings (e.g.,
Chamaecyparis, Thuja, Thujopsis, Cupressus, and Neocalli-
tropsis). The wings may be fully reduced, rarely in a form of
narrow longitudinal rims (Platycladus, Microbiota, and Jun-
iperus), or subapically extended (Calocedrus). Wings of Fok-
ienia seeds are rather unequal, spreading, and variable in size.
Seeds of some of the Southern Hemisphere genera are distin-
guished by the presence of three wings (Fitzroya and Actino-
strobus), and others have a single main wing, with a second
KVAC

EK ET AL.CUPRESSACEAE IN THE NORTH AMERICAN TERTIARY 339

Fig. 5 Cones, seeds, and foliage of Tetraclinis brachyodon from the European Tertiary for comparison with Tetraclinis salicornioides. A,
Seed cone on twig showing subcentral umbo, small seed cone on twig with subcentral umbos, St. Zacharie, Oligocene; MNHN ZK2, #2.5. B,
Small cone with subterminal umbos, St. Zacharie, Oligocene; MNHN ZK3, #2.5. C, One cone of several attached to a twig, Radoboj, Middle
Miocene (orig. Kovar-Eder and Kvacek 1995); Univ. Graz 11424, #2.5. D, Larger cone, Armissan, Upper Oligocene; MNHN 11155, #2.5.
E, Lectotype of T. brachyodon, ParisMont Rouge/Chatillon, Middle Eocene (orig. Brongniart 1822, pl. 5, g. 3A); MNHN 78, #2.5. F, Twig
showing alternate branching typical of this species; MNHN s.n., #1. G, Widely opened cone, Aix-en-Provence, Oligocene-Miocene boundary;
MNHN s.n., #2.5. H, Seed with deeply cordate base, St. Zacharie; MNHN ZK8, #2.5. I, Seed with very truncate base, Armissan, Upper
Oligocene (orig. Saporta 1862, pl. 1, g. 6C); MNHN 11157, #2.5. J, Twig with two terminal male cones (?), Oligocene, St. Zacharie; MNHN
16320, #2.5. K, Slender twig, Arcueil, Middle Eocene (orig. Watelet 1866, pl. 32, g. 3); MNHN 7828, #2.5. L, Detail from g. 5F, #2.5.
M, Abaxial cuticle of the specimen shown in g. 5F and 5L, #250.
wing that is much reduced (Libocedrus, Austrocedrus, and
Pilgerodendron). The most common type of seed has two
wings that may form only a narrow longitudinal rim (e.g.,
Chamaecyparis, Thuja, Thujopsis, Cupressus, and Neocalli-
tropsis). The wings may be fully reduced (Platycladus, Micro-
biota, and Juniperus) or apically enlarged and fused (Calo-
cedrus). Wings of Fokienia seeds are rather unequal, spreading,
and variable in size. Seeds of some of the Southern Hemisphere
genera deviate by the presence of three wings (Fitzroya and
Actinostrobus), and others have only a single large lateral
wing, with the other wings being markedly reduced or absent
(Libocedrus, Austrocedrus, and Pilgerodendron). In Tetra-
clinis, the seeds are principally biwinged, with wings expand-
ing basally (e.g., g. 1L, 1M), but the two innermost seeds of
340 INTERNATIONAL JOURNAL OF PLANT SCIENCES
the cone bear a third wing in the form of a slight medial keel.
The unique form of Tetraclinis seeds among the Cupressoids
provides an important character for the identication of fossil
representatives.
Foliage of Cupressaceae
The Cupressoids may have either decussate or verticillate
phyllotaxy, but the foliar morphology is variable in terms of
many features. Pronounced variation occurs during ontogeny,
such that acicular leaves (free or partly decurrent) are typically
developed in young stages but survive only exceptionally on
adult plants (e.g., Juniperus subg. Juniperus). The acicular fo-
liage type tends to make transitions through ontogeny to the
scale leaves that prevail in the adult foliage of most genera.
Even in mature foliage, there may be considerable foliar var-
iation according to position within the shoot system. As a rule,
scale leaves of the stalks of seed cones tend to be reduced,
shortened, and radially disposed. Branches in cross section may
be either biconvex (tetragonal; octagonal in Neocallitropsis),
with more or less homomorphic leaves (e.g., Cupressus and
Fitzroya), or more or less attened (to convex-concave) and
either more or less homomorphic (e.g., Pilgerodendron and
Thujopsis) or with dimorphic leaves (e.g., Fokienia). Even epi-
dermal characteristics are of limited value for generic distinc-
tions within this family (Florin 1931). Therefore, botanists
dealing with this group of conifers generally prefer the char-
acters of seed cones for classication of the family and for
distinction of genera (e.g., Li 1953; Gaussen 1968).
In the Cupressaceae, foliage morphology can vary among
species of the same genus (Calocedrus decurrens vs. Calocedrus
macrolepis, Calocedrus formosana). Conversely, multiple gen-
era may share the same type of foliage (Chamaecyparis and
Thuja; Cupressus and Juniperus). Branching patterns are also
variable within a genus or even a species (Rouane 1973). Op-
posite branching is characteristic of some Southern Hemi-
sphere Cupressoids (Libocedrus and Austrocedrus), but it oc-
curs occasionally in other genera. Opposite branching was
used as an argument by Heer (1855, 1870) and others to assign
fossil sterile branches to Libocedrus. However, studies of fossil
foliage with associated or attached cones have revealed that
opposite branching was widespread in various genera of an-
cient Cupressoids (McIver 1994).
Leaves of extant Cupressaceae are never fully fused to form
cladode-like segments; however, this condition is now known
in two fossil taxa: Tetraclinis salicornioides and Fokieniopsis
catenulata (Bell) McIver et Basinger. Both species are similar
in branching architecture and in the presence of fused foliar
segments. They differ in epidermal anatomy and cone mor-
phology. It might be tempting to use the specialized feature of
fully fused, cladode-like segments to relate both taxa. However,
this may be a case of convergent or parallel evolution, as is
typical of both extant (Chamaecyparis and Thuja) and extinct
(Mesocyparis and Chamaecyparis corpulenta) members of this
family.
Most Cupressoids show distinct Florin rings around the sto-
matal cavity and often also show distal papillae on ordinary
cells (Oladele 1983). These characters help distinguish the Cu-
pressaceae from the Cheirolepidiaceae, which shows papillae
overlapping the stomatal pits. Distinct Florin rings and papillae
on ordinary cells are clearly developed in both European and
North American records of fossil Tetraclinis foliage. Distri-
bution and arrangement of stomata provide additional diag-
nostic characters, but they are not as reliable since they are
inuenced by exposure in light (Fitting 1942). Kvacek (1989)
reasoned that the differences in stomatal topography between
the species of Tetraclinis probably arose as a result of ecolog-
ical factors, such as adaptation to light conditions and hu-
midity. In Tetraclinis brachyodon, which occurs earlier in the
fossil record (Early Eocene to Early Pliocene), the foliage is
only slightly heteromorphic, and the stomata are conned to
the adaxial surface of the leaf and the area adjacent to the leaf
margin. In contrast, the stomata of T. salicornioides (Middle
Eocene to Early Pliocene) are scattered all over the surface of
segments (see Friis 1977), which indicates conditions of diffuse
light in closed forests. Tetraclinis brachyodon and T. salicor-
nioides commonly occur together, although in indirect pro-
portion. Tetraclinis brachyodon grew abundantly in more
southerly localities in the European Tertiary (Aix-en-Provence,
Armissan, St. Zacharie, Ha ring, Kumi, and Radoboj), which
are associated with subhumid forest vegetation (open semi-
evergreen sclerophyllous woodland sensu Mai [1994]). The
xeromorphic character of the genus has become most pro-
nounced in the Mediterranean (summer-dry) type of climate,
where Tetraclinis articulata occurs today.
Review of Other Fossil Cupressaceae in Relation
to Tetraclinis salicornioides
Putative Cupressoids have been reported from the Jurassic
and Early Cretaceous, but there is no convincing evidence of
this group until the Late Cretaceous. Many of the fossils with
decussate scale leaves similar to those of the Cupressoids can
be distinguished as members of the extinct Cheirolepidiaceae,
because the cuticle reveals a papillate rim overlapping the sto-
matal pit. Examples include Cupressinocladus ramonensis
Chaloner et Lorch (1960), Cupressinocladus micromerum
(Heer) Pais (1974), and Paleocyparis exuosa Saporta et Mar-
ion teste Neves (1950). A comprehensive revision of such fos-
sils is needed to elucidate the earliest records of true Cupres-
saceae. One of the Cretaceous fossils formerly assigned to
Chamacyparis is Chamacyparis cretacea Velenovsky et Vinik-
la r fromthe Cenomanian of Bohemia (Velenovsky and Vinikla r
1926). Inspection of the type specimens revealed helically ar-
ranged needle-like leaves and associated Sequoia-like cones (Z.
Kvacek, personal observation). Libocedrus salicornioides cre-
tacea Velenovsky (1885) represents a poorly preserved cone
scale of Dammara borealis Heer (Kvacek 1989).
In his classication of the Cupressaceae and Taxodiaceae,
which were merged into one family, Eckenwalder (1976) ex-
pects primitive traits of the cupressoid group from the
Northern Hemisphere: globular, many-scaled seed cones; cru-
ciate, terete branchlets; and arborescent habit. He considers
Cupressus to be most similar to the probable ancestor of the
tribe Cupresseae, from which some members of his subtribe
4, namely Tetraclinis, Widdringtonia, Libocedrus, and Aus-
trocedrus, could have originated ) by reduction of the two
pairs of ovuliferous cone scales (Eckenwalder 1976, p. 252).
In the view of McIver and Aulenback (1994), however, the
earliest known cupressoid seed cones fromthe Cretaceous bore
KVAC

EK ET AL.CUPRESSACEAE IN THE NORTH AMERICAN TERTIARY 341

four scales, thus differing markedly from the many-scaled Cu-
pressus-like cones of the hypothetical ancestor. The rst record
of Cupressus-like cones with many scales was not found until
the late Paleocene or early Eocene (Cupressoconus machenryi
(Baily) Boulter et Kvacek [1989] from Ireland). More studies
on the cones of Cretaceous conifers are needed to trace the
history of the Cupressaceae. In this respect, Widdringtonites
Endlicher (W. reichii [Ettingsh.] Heer) and Tetraclinopsis Boyd
(T. paututensis Boyd), which have noncolumellate cones with
4 [5] woody valvate cone scales (see Boyd 1992) but helically
(?) disposed scale leaves on twigs deserve attention.
Among the Upper Cretaceous foliage with decussate phyl-
lotaxy, those showing Florin rings around the stomatal pits
most likely belong to the Cupressoids. Some of these shoots
bear cones. The most common fossils of this kind have been
assigned to Thuja cretacea (Heer) Newberry, were distributed
at high latitudes of the Northern Hemisphere (Samylina 1988),
and ranged into the Early Tertiary (Thuja ehrenswaerdii [Heer]
Schweitzer, Thuja polaris McIver and Basinger). Their cones
have imbricate scales that are similar in most respects to those
of extant Thuja species (McIver and Basinger 1989). Meso-
cyparis (McIver and Basinger 1987; McIver and Aulenback
1994; syn. Microconium Golovneva 1988, 1994) differs from
the extant Cupressoids in that it bears cones in decussate or
distichous pairs. Its globular quadrivalvate and columellate
cones resemble those of Chamaecyparis nootkaensis (D. Don)
Spach in possessing four more or less equal cone scales (McIver
and Aulenback 1994). A true Chamaecyparis, with minute
cones of four nearly equal scales, was described from the Late
Cretaceous of Canada (McIver 1994). None of these taxa has
a foliage with cladode-like segments.
Another species with cladode-like foliar branches resembling
those of Tetraclinis salicornioides occurs in the Cretaceous and
Paleocene and has gone under the names Androvettia caten-
ulata Bell, Ditaxocladus planiphyllus Guo et Sun, Fokienia
catenulata (Bell) Brown, Fokienia ravenscragensis McIver et
Basinger, Fokieniopsis catenulata (Bell) McIver et Basinger, and
Libocedrus catenulata (Bell) Kryshtofovich. It is known from
Canada (Bell 1949; Brown 1962; McIver and Basinger 1990)
and from central and eastern Asia (Kryshtofovich and Bai-
kovskaya 1966; Romanova 1975; Guo et al. 1984; Samylina
1988). Subglobose seed cones with eight to 10 woody scales
were attached to twigs of this foliage in opposite pairs, as was
observed in specimens from Saskatchewan (McIver 1992) and
Xinjiang (S. R. Manchester and S.-X. Guo, unpublished data,
1995). Additional comparative work and careful attention to
the nomenclatural rules of priority are needed to verify the
appropriate binomial for this species, but for purposes of dis-
cussion in this article, we refer to it as F. catenulata (Bell)
McIver et Basinger. The seed cones differ from the those of
extant Fokienia Henry et Thomas based on their scales, which
are valvate basally and spatulate to subpeltate apically. The
seeds of this plant remain uncertain. In some cases, the as-
sociated seeds illustrated by McIver (1992) show a spinulose
wing margina character not known in extant coniferous
seeds.
Although F. catenulata resembles T. salicornioides by its
cladode-like segments, the sterile foliage of the two taxa may
be distinguished by subtle, partly overlapping differences: in
F. catenulata, the branch systems are more slender in outline,
the veins to lateral branches often arise well above the base
of the medial leaf segments (Kryshtofovich and Baikovskaya
1966, pl. 6, g. 3; Guo et al. 1984, pl. 1, g. 5a; McIver
1992, text, g. 1), and the epidermis is nonpapillate and
shows straight-walled cells and short rows and groups of
stomata with only narrow Florin rings. In T. salicornioides,
the branch system is usually fan shaped (Engelhardt 1886;
Ferguson 1971), the pair of vascular bundles leading to the
lateral branches arises more often at the base of the leaf
segment (g. 2I2L), the cuticle is mostly papillate, the cell
walls are usually undulate with thickenings, and the stomata
are arranged in longer rows within several longitudinal sto-
matal areas. Stomata bear thick Florin rings. In extant Fok-
ienia, the foliage is not fused or cladode-like, and the scale
leaves have more densely spaced stomata that occur in nar-
row, well-demarcated bands (Florin 1931, pp. 438440; Z.
Kvac ek, personal observation).
We studied cuticle from leaves of F. catenulata (Bell) McIver
et Basinger from the Paleocene Fort Union Formation of Sand
Draw, Wyoming (UF locality 18131) for comparison with the
T. salicornioides fossils. The specimens examined included a
part of the ultimate branch and a large medial segment. Cuticle
of the cladode-like segments in F. catenulata is much thinner
than that of T. salicornioides and lacks papillae. The ordinary
epidermal cells are straight walled to very slightly and nely
undulate, polygonal-quadrangular, and usually isodiametric or
up to three times longer than wide (width 1525 mm, length
2550 mm), becoming narrower and longer on the edges of
the cladode-like segment. The monocyclic stomata form short
longitudinal and irregular groups in stomatal areas. The sto-
mata are irregularly oriented with apertures that are (12) 20
mm wide and (17) 25 (30) mm long and are bordered by a
thin Florin ring.
The foliar similarities between F. catenulata and T. salicor-
nioides can be explained by parallel evolution because their
cones are different. The attached cones of Fokieniopsis, called
Fokienia ravenscragensis by McIver et Basinger (1990), have
twice as many decussate scales and are borne in opposite pairs.
The Tetraclinis fossils have four-scaled cones borne singly or
in pairs on the twigs in T. salicornioides (Ferguson 1971; Kva-
cek and Walther 1995) or singly in T. brachyodon (Kovar-Eder
and Kvacek 1995). Basal cone scales are more or less cordate,
and their form is reected in the shape of the winged seeds.
Biogeographic Considerations
In most systems of conifers, Tetraclinis is classied close to
the group of Afro-Australian genera of the Cupressaceae (Wid-
dringtonia Endl., Actinostrobus Miq. ex Lehm., and Callitris
Vent.), although it stands out amongst these as perhaps much
less close to the others (Page 1990, p. 305). Although some
authors have favored a Southern Hemisphere origin for Te-
traclinis (Hart 1987), the fossil record demonstrates that the
genus had a substantial distribution in the Tertiary of the
Northern Hemisphere. Mai (1994, 1997, 1998) indicated that
Tetraclinis brachyodon and T. salicornioides were not clearly
distinct species. However, no brachyodon-like foliage has been
recovered so far in the North American localities of Tetraclinis.
Tetraclinis brachyodon tends to have more attened foliar
342 INTERNATIONAL JOURNAL OF PLANT SCIENCES
shoots, up to three male cones (Saporta 1862), and larger
umbos (Gaussen 1968).
The distribution of T. salicornioides indicates that the species
must have migrated between western North America and Eu-
rope during the early or middle Tertiary. Although it extended
through Europe to the Transcaucasus area (Kvacek 1989; Mai
1994, 1995; Kovar-Eder et al. 1996), the absence of Tetraclinis
from localities east of the Caspian Sea indicates that the genus
did not cross Asia. (Z. Kvacek examined the specimen iden-
tied as Libocedrus salicornioides from core 138 near the
Krugloe Sea, Kazakhstan [Zhilin 1974, p. 97] and observed
that the twig fragment does not consist of cladode-like seg-
ments and corresponds most likely to Chamaecyparis or
Thuja.) Therefore, we infer that T. salicornioides communi-
cated directly between Europe and North America, apparently
across the North Atlantic. Based upon shared occurrences in
the early Tertiary of North America and Europe, many genera
of angiosperms are inferred to have crossed the North Atlantic
prior to the rifting apart of Canada, Greenland, Iceland, and
Scandinavia (Tiffney 1985; Manchester 1999). In North Amer-
ica, T. salicornioides has not been recognized prior to the early
Oligocene, and it seems to be restricted to the Pacic North-
west (Oregon, Washington, and Idaho), where it persisted at
least to the middle Miocene. The species was more widespread
in Europe, ranging from southern France (Saporta 1865) to
Transcaucasia (Kolakovskij and Shakryl 1978, pl. 1, gs. 3,
4) and from the middle Eocene to the early Pliocene. Both the
North American and European varieties with mesophytically
adapted foliage became extinct by the end of the Tertiary, but
the T. brachyodon type persisted in southern Europe, with the
closely similar extant species Tetraclinis articulata persisting
today in the western Mediterranean.
Many of the genera shared between Europe and North
America during the early Tertiary are now found to survive in
Asia (Tiffney 1985; Manchester 1994). However, Tetraclinis,
which is lacking in Asia today and which is unknown in the
Tertiary oras of eastern Asia, does not conform to this pat-
tern. At the time during which Tetraclinis was spreading be-
tween Europe and North America, the Turgai seaway effec-
tively blocked many plants from crossing between Europe and
Asia. Many other plants succeeded in traversing the Turgai
region as the seaway receded in the Oligocene (see Kvacek
1994; Tiffney 1994); however, for unknown reasons, Tetra-
clinis remained absent from eastern Asia.
Acknowledgments
Specimens were made available by D. Erwin, W. C. Rember,
C. J. Smiley, Scott Wing, Dario De Franceschi, and other keep-
ers of the herbaria and fossil collections. Helpful comments
were provided by D. H. Mai, E. E. McIver, and H. W. Meyer.
Manuscript review comments were kindly provided by James
Eckenwalder and an anonymous reviewer. This research was
supported cooperatively by Czech Ministry of Education grant
ME 054 to Z. Kvacek and National Science Foundation grant
INT 560260112 to S. R. Manchester.
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