Int. J. Plant Sci. 169(7):918–927. 2008.

Ó 2008 by The University of Chicago. All rights reserved.

1058-5893/2008/16907-0009$15.00 DOI: 10.1086/589694

WHOLE-PLANT RECONSTRUCTIONS IN FOSSIL ANGIOSPERM RESEARCH

Zlatko Kvaček1
Charles University in Prague, Faculty of Science, Albertov 6, CZ-128 43 Prague 2, Czech Republic

There have been many recent advances in angiosperm paleobotany based on studies of reproductive organs,
but it remains important that ‘‘whole-plant’’ concepts not be neglected. Reconstructions of whole fossil plants
based on foliage, fruits, seeds, flowers, pollen in situ, and wood are not always easy, but where done well they
often yield important new insights. The most convincing whole-plant reconstructions are accomplished when
organs that are normally found separately are found attached. Such occurrences are relatively rare. The repeated
co-occurrence of dispersed organs that share anatomical features provides another acceptable line of evidence for
reconstructions. However, mere co-occurrence, particularly in the case of extinct lineages, provides less-
convincing evidence. An important way to connect dispersed pollen and macrofossils is the study of pollen in situ.
Several examples from Platanaceae, Fagaceae, Cercidiphyllaceae, Malvaceae s.l., and Cucurbitaceae show the
importance of whole-plant approaches for evolutionary and other interpretations.

Keywords: Cucurbitaceae, Fagaceae, fossil angiosperms, whole-plant reconstructions, Tertiary, Malvaceae s.l.,
Platanaceae.

Introduction in Wetterau. However, this reconstruction included in part

One major difficulty in studies of fossil angiosperms and other
plant groups is that the fossil record consists mainly of plant frag-
ments. By contrast with living plants, which are usually collected
as flowering or fruiting specimens of whole plants or their parts
(e.g., twigs), in fossil plants, individual organs are detached in
most cases. From the earliest days of paleobotanical research,
scientists dealing with such fossils have attempted to combine
knowledge, at least from foliage and fruits, to get a more com-
plete reconstruction of the original living plant.
Early protagonists of this holistic concept in studies of fossil
angiosperms were Franz Unger (1800–1870), Austrian botanist
and professor of botany at Graz and Vienna, and Oswald Heer
(1809–1883), Swiss paleontologist and professor of paleontology
at Zürich. Some of their reconstructions of fossil angiosperms
are partly correct and still remain valid. Unger, for example, was
successful in a reconstruction of Paliurus favonii Ung. based
both on fruits and foliage from the Miocene of Parschlug (Unger
1847; see also Bůžek 1971). Similarly, the reconstruction of
Podogonium Heer, which Heer (1859) considered an extinct
legume, was correct. Gregor and Hantke (1980) challenged
this interpretation and transferred Podogonium knorrii Heer to
Gleditsia, but after more-detailed comparison with modern le-
gumes, Herendeen (1992) confirmed Heer’s original view. This
whole-plant reconstruction and its relationship to Podogonium
are also supported by more-complete specimens and a study of
detached flowers with pollen in situ (Mohr and Gregor 1984;
Liu et al. 2001), as well as by the same association of fossil fruits
and foliage in China and Japan as in Europe (Wang et al. 2005).
Although some early attempts at whole-plant reconstruc-
tions were successful, others turned out to be misleading. Un-
ger (1860), for instance, believed that he had reconstructed the
plant of Annona (Annonaceae) from the Miocene of Salzhausen
1
E-mail: kvacek@natur.cuni.cz.
Manuscript received February 2008; revised manuscript received April 2008.
seeds of Magnolia (Mai 1975) and leaves belonging to Nyssa (Z.
Kvaček, unpublished observations). Another of Unger’s errors
was interpreting detached fruit valves as elm samaras (Unger
1847, as Ulmus bronnii). These common fossils have since
been attributed to an unrelated genus, Craigia (Kvaček et al.
1991), and are discussed in more detail in ‘‘Malvaceae s.l.’’
Like Unger, Heer (1856, 1859) also misinterpreted certain fos-
sil angiosperms as he attempted reconstructions. For example,
from leaf venation, Heer believed that one of the commonest
leaf fossils in the Miocene of Oehningen, superficially similar to
Tilia, actually belonged to Ficus. He supported this view with a
fossil fruit that, according to Kirchheimer (1957), was too poorly
preserved to have taxonomic value. Epidermal studies have sub-
sequently suggested (Knobloch and Kvaček 1965a, 1965b; Wo-
robiec 2003), that such ‘‘Ficus’’ leaves, at least those occurring
in the Neogene of Europe, are those of a malvalean plant and
are properly known as Byttneriophyllum Givulescu ex Knobloch
& Kvaček (1965b). Heer (1856) also wrongly combined two
other common plant elements from the European Tertiary and
assigned the resulting reconstruction to Grewia (Malvaceae s.l.).
Brown (1935) later recognized Cercidiphyllum (Cercidiphylla-
ceae) among these characteristic leaves. The fruits, which Heer
believed to belong to the same plant, were reidentified by Kirch-
heimer (1957) as endocarps of Celtis (Celtidaceae).
The failure of many previous reconstructions of fossil angio-
sperms, combined with the realities of the plant fossil record,
has inevitably led to independent studies of detached organs. In
particular, studies of fossil fruits, seeds, and flowers preserved as
carbonized mesofossils have achieved much success and have
contributed substantially to our current understanding of angio-
sperm evolutionary history (Friis 1983; Friis et al. 1992, 1994,
among many others). Similar progress has also been made in pa-
leopalynology, particularly in using SEM and TEM observations,
as well as in studies of wood anatomy and epidermal anatomy.
However, notwithstanding all of this progress, it is also impor-
tant that the whole plant concept not be neglected.

918
 KVAČEK—WHOLE-PLANT RECONSTRUCTIONS OF FOSSIL ANGIOSPERMS
Revival of the Whole-Plant Concept
Revival of a holistic approach to reconstructing whole fossil
angiosperms was initiated by those searching for more-complete
fertile specimens in which reproductive structures and leaves are
attached. Such discoveries are rare but provide convincing evi-
dence about the interconnection of otherwise detached organs.
Plant fossils of Cretaceous age that show these connections
include those assigned to Prisca Rettalack & Dilcher (1981),
Archaefructus Sun et al. (1998, 2002), among others. Examples
from the Paleogene and Neogene include, for example, extinct
Fagopsis Hollick (Manchester and Crane 1983), Cedrelospermum
Saporta (Manchester 1987, 1989), monocotyledonous Limno-
biophyllum Krassilov (Stockey et al. 1997), and an aquatic plant
assigned to Decodon (Kvaček and Sakala 1999). However, how
far the affinities of these important specimens are understood
depends greatly on the quality of preservation. Although it is ad-
visable to use all available techniques to ascertain characteristics
of pollen in situ, cuticles, and details of the fructifications, this is
not always possible. The key rare specimens that show attach-
ment between different organs are often preserved as impres-
sions (e.g., extinct Pseudofagus Smiley & Huggins [Smiley and
Huggins 1981] or Pseudosalix [Boucher et al. 2003]).
Beyond these relatively rare examples of fertile plant speci-
mens, we are still at a loss to demonstrate the connections among
most of the plant fossils that we encounter. Nevertheless, several
lines of argumentation have been employed to infer connections
between detached fossil plant organs never found in organic con-
nection: (i) repeated co-occurrence; (ii) identical or similar epider-
mal anatomy; (iii) identical or similar systematic affinities; (iv)
pollen in situ and pollen ex situ (accumulations scraped from the
surface of leaves). In some cases, one or more of these arguments
have been combined to justify a reconstruction.
The following examples illustrate some of the ways in
which reconstructions of Tertiary angiosperms have been as-
sembled. Most of the plants comes from the Tertiary of Cen-
tral Europe, in particular from the Bohemian Massif and
adjacent areas. Details of research methods, occurrences, and
other circumstances of elements discussed are cited in the re-
spective publications and are not repeated here.
Platanaceae
The Tertiary platanoid Platanus neptuni (Ettingsh.) Bůžek,
Holý & Kvaček is well known from previous studies based on
its simple, unlobed leaves (Bůžek et al. 1967; Knobloch 1973;
Hably 1980; Walther 1985). It is also known from infructescen-
ces recalling heads of Sparganium or even Liquidambar (fig.
1A), as well as by staminate inflorescences (fig. 1C) with pollen
in situ (fig. 1D) and vegetative organs such as foliage branches
(fig. 1B), isolated leaves, and stipules. All organs have been
combined into a single species because of the shared presence
of the same type of glandular trichomes (fig. 1E) and the regu-
lar co-occurrence of detached leaves, stipules, inflorescences, in-
fructescences, fruitlets, and stamens at Markvartice in the
Oligocene of northern Bohemia (Bůžek et al. 1967) and else-
where. The plant is extinct and differs from living Platanus in
the manner of attachment of the single head to the stalk, which
is demarcated below by a narrow circuital rim (Kvaček and
919
Manchester 2004). This ancient plane tree is not related to the
Asian Platanus kerrii Gagnep. (subgenus Castaneophyllum Le-
roy), which has also simple, unlobed but evergreen foliage. Plat-
anus kerrii differs from P. neptuni and all other living plane
trees in that the petiole does not cover the bud. Platanus neptuni
has been separated into an extinct subgenus Glandulosa Kvaček,
Manchester & Guo (Kvaček et al. 2001) because of an unusual
attachment of single heads on stalks and peculiar epidermal
anatomy. In most cases, the foliage is simple and lanceolate,
but compound trifoliolate and quinquefoliolate leaves are also
known. They all share the same epidermal pattern. For prag-
matic reasons, the detached organs of P. neptuni were assigned
to several morphoforms (Kvaček and Manchester 2004).
Fossils of P. neptuni are often found in volcanic areas where
permineralized wood of the Platanus type is also present. The
wood is characterized by broad homocellular rays and scalari-
form perforation plates with many bars. According to Sakala
(2006), this type of wood may represent trunks or branches of
P. neptuni. More studies are required to better understand vari-
ation in fossil Platanus-like wood. Currently, wood of this kind
is split into several narrow morphotaxa.
Fagaceae
Contrary to the situation with Platanus, some common faga-
ceous fossils that typically occur in the Paleogene of Europe
have been assigned not to a modern genus but to the extinct ge-
nus Eotrigonobalanus Walther & Kvaček (Kvaček and Walther
1989; Walther 2000). In this case, the fruits were originally
assigned to the living genus Trigonobalanus s.l. (Mai 1970). At
present, living trigonobalanoids are split into three monotypic
genera (Trigonobalanus s.s., Colombobalanus, and Formano-
dendron), and in this respect, separation of the fossils into an
independent extinct genus appears to be appropriate. The fos-
sils differ in many ways from all three living genera.
In Eotrigonobalanus, the foliage (fig. 2A, 2B) and fruits (fig.
2D, 2E), which are found occasionally in catkinlike infructescen-
ces (Kvaček and Hably 1998), are treated separately as two mor-
phospecies, Eotrigonobalanus andreanszkyii (Mai) Walther &
Kvaček for fruits and Eotrigonobalanus furcinervis (Rossm.)
Walther & Kvaček for leaves. This is a common approach when
naming detached foliage and reproductive organs in many other
fossil angiosperms. Epidermal anatomy of E. furcinervis (fig. 2F)
is very characteristic and sufficient to allow identification of leaf
fragments. Staminate catkins of Eotrigonobalanus have not yet
been recovered, and pollen in situ remains unknown. Only the
regular co-occurrence of dispersed pollen of the Fususpollenites
type, also called Eotrigonobalanus eissmannii Walther & Zetter
(Walther and Zetter 1999), suggests that this morphotype (fig.
2C) may belong to Eotrigonobalanus. This pollen, with faga-
ceous morphology, but not corresponding to any particular ex-
tant form, was found in concentrated masses on the leaf surfaces
of Eotrigonobalanus.
In the Evros Mountains, Thrace, northeastern Greece, accumu-
lations of Eotrigonobalanus foliage accompany permineralized
wood of the evergreen Quercus type, which may represent trunks
of Eotrigonobalanus (Velitzelos et al. 1999). Several morphotaxa
of Eotrigonobalanus wood have been described. According to Sel-
meier and Velitzelos (2000), this wood requires more comparison
920 INTERNATIONAL JOURNAL OF PLANT SCIENCES

Fig. 1 Platanus neptuni (Ettingsh.) Bůžek, Holý & Kvaček. A, Immature infructescence, lectotype, Kuclı́n, northern Bohemia, Eocene, 34. B, Leafy
twig, Kuclı́n, northern Bohemia, Eocene, 31. C, Staminate inflorescence, Markvartic, northern Bohemia, Oligocene, 35.5. D, Pollen in situ (SEM), Plesná,
core V 146, western Bohemia, Oligocene, 31500. E, Abaxial cuticle of a leaf compression, Plesná, core V 146, western Bohemia, Oligocene, 3350.

with the natural variation in wood anatomy seen in comparable
tropical Fagaceae.
Malvaceae s.l.
The case of fossil Craigia exemplifies a situation in which the
name of the living genus is applied to reproductive organs, while
various morphogenera are used to accommodate vegetative re-
mains. Two extinct whole-plant species can be differentiated in
the Northern Hemisphere (Kvaček et al. 2005). They are hardly
distinguishable in fruit morphology but differ in the accom-
panying foliage type. Craigia bronnii (Ung.) Kvaček, Bůžek &
Manchester from Europe (and Spitsbergen) is more completely
known. In addition to dispersed fruit valves, which are common,
complete fruits are also known in various stages of maturity (fig.
3E), with seeds in situ. Dispersed flowers and flower buds (fig.
3D) with pollen in situ (fig. 3C) have also been recovered (Pingen
et al. 2001; Kvaček et al. 2002; Kvaček 2005).
The North American and Asian Craigia oregonensis (Arnold)
Kvaček, Bůžek & Manchester is known mostly from detached
fruit valves. However, in both locations, the pollen in situ or the
co-occurring dispersed pollen is of the Intratriporopollenites (i.e.,
tilioid) type. Kvaček et al. (2005) hesitated to assign the foliage
accompanying fruits to the living genus and left both entities un-
der traditional morphotaxa: Dombeyopsis lobata Ung. and Plaf-
keria obliquifolia (Chaney) Wolfe for the North American and
Asian specimens, respectively. The two modern species of the ge-
nus also differ from each other more in foliar characters than in
fruit morphology. Only the large, mostly trilobate leaf fossils
from Europe (fig. 3A, 3B) have yielded epidermal features that
match those of Craigia, but these features are also found in other
malvalean genera (Kvaček 2005).
Recent studies (Sakala 2007) revealed a new type of fossil mal-
valean wood, probably belonging to Craigia in the Oligocene of
 KVAČEK—WHOLE-PLANT RECONSTRUCTIONS OF FOSSIL ANGIOSPERMS 921

Fig. 2 Eotrigonobalanus furcinervis (Rossm.) Walther & Kvaček/Eotrigonobalanus andreanszkyi (Mai) Walther & Kvaček/Eotrigonobalanus
eissmannii Walther & Zetter. A, Leaf impression, Staré Sedlo, western Bohemia, Eocene, 31.3. B, Leaf compression, Haselbach, Saxony,
Oligocene, 31.1. C, Pollen from the leaf surface (SEM), Zeitz, Saxony, Eocene, 34000. D, Fruit cupule, Etzdorf, Saxony, Eocene, 34.5. E,
Specimen in D from reverse side, 34.5. F, Abaxial leaf cuticle, Witznitz, Saxony, Oligocene, 3380.

northern Bohemia. The wood is semi–ring porous with vessels
mostly solitary or in short, radial multiples. Again, the problem
of variation in wood structure of this type requires further com-
parisons with extant taxa. The situation is further complicated
by the fact that other malvalean wood types occur with Craigia
fossils in volcanic areas of the Czech Republic, for example, co-
occurring fossil species of Tilia known by leaves and fruiting
bracts (Kvaček and Walther 2004). Studies by Manchester
et al. (2006), who characterized the wood of extant Craigia,
have facilitated further comparisons.
Cercidiphyllaceae
Remains of Cercidiphyllum, which are widely distributed in
the European Tertiary, were revised by Jähnichen et al. (1980),
who recognized two morphospecies: Cercidiphyllum crenatum
(Ung.) R. Brown for leaves and Cercidiphyllum helveticum
(Heer) Jähnichen, Mai & Walther for fruits. Later studies of
Tertiary plants from the Czech Republic brought surprising
new information (Kvaček and Konzalová 1996). In particular,
the discovery of fertile branches with fruits attached (Oligocene
of Bechlejovice; fig. 4B), matching living Cercidiphyllum japo-
nicum Sieb. & Zucc., made it unnecessary to assign fruits to an
independent morphospecies.
One fossiliferous layer within the well-known Bı́lina Mine
in northern Bohemia, of Early Miocene age, also yielded stami-
nate inflorescences in various stages of maturity in association
with the leaves and fruits of C. crenatum. While undeveloped
inflorescences appear similar to those of the living Cercidiphyl-
lum (fig. 4D), fully mature inflorescences are different, with
922 INTERNATIONAL JOURNAL OF PLANT SCIENCES

Fig. 3 Dombeyopsis lobata Ung./Craigia bronnii (Ung.) Kvaček, Bůžek & Manchester. A, Leaf, Bı́lina Mine, N. Bohemia, Miocene, 31. B, Leaf,
lectotype, Brestany, northern Bohemia, Miocene, 31.3. C, Pollen in situ (SEM), Hambach, western Germany, Miocene, 31700. D, Flower bud,
Hambach, western Germany, Miocene, 320. E, Complete capsule, Bı́lina Mine, northern Bohemia, Miocene, 33.5.

long-stalked flowers arising from the basal scales in groups
of about four. Each flower has a pentamerous perianth (fig.
4G) and several stamens with long filaments (fig. 4C). Thus,
this fossil Cercidiphyllum is clearly not identical with the liv-
ing species, which has naked male flowers. It does, however,
provide evidence of the likely primitive condition in the group
and supports the inclusion of Cercidiphyllaceae in Saxifra-
gales, as emphasized by molecular data (Fishbein et al. 2001).
In all other details, including the morphology of stamens and
pollen in situ (fig. 4A), as well as leaves (fig. 4E) and fruits
(fig. 4F), C. crenatum matches living Cercidiphyllum. The re-
duction of staminate inflorescences in living representatives
can be explained as paedomorphic. Living Cercidiphyllum is
arrested in the development of staminate flowers in a juvenile
stage.
Fossil wood of the Cercidiphyllum type co-occurs with C.
crenatum in volcanic areas in northern Bohemia. It is assigned to
Cercidiphylloxylon kadanense Prakash et al. (1971). This species
of fossil wood shares many features with living Cercidiphyllum,
according to Sakala and Privé-Gill (2004): diffuse-porous wood
with mostly solitary vessels, angular in outline; scalariform perfo-
ration plates with ;40 bars; and heterocellular, mostly bi- to
triseriate rays. This plant provides the oldest record of wood at-
tributable directly to the genus Cercidiphyllum.
 KVAČEK—WHOLE-PLANT RECONSTRUCTIONS OF FOSSIL ANGIOSPERMS 923

Fig. 4 Cercidiphyllum crenatum (Ung.) R. Brown. A, Pollen in situ, Bı́lina Mine, northern Bohemia, Early Miocene, 3600. B, Fertile leafy twig (position
of fruits marked by arrows), Bechlejovice, northern Bohemia, Oligocene, 31. C, Staminate inflorescence with long-stalked flowers, showing narrow perianth
and stamens, Bı́lina Mine, northern Bohemia, Miocene, 32.2. D, Pair of immature staminate inflorescences, same locality as C, 32. E, Leaf impression, same
locality as C, 31. F, Group of fruits, same locality as C, 33. G, Staminate inflorescence with pentamerous perianth devoid of stamens, same locality as C, 33.

Nyssaceae
Nyssa bilinica (Ung.) Kvaček is another example of a more
completely known fossil plant that does not match any single
living representative of the genus. It was recovered in the Bı́lina
area, together with Cercidiphyllum and other plants from the
swamp assemblage preserved in the clay just above the worked
coal seam.
Foliage of N. bilinica is well characterized by epidermal anat-
omy with spatulate trichomes (fig. 5C; see also Kvaček and Bůžek
1972). Two leaf forms can be recognized: large, more mesomor-
phic shade leaves with visible stomata and less distinct papillae on
924 INTERNATIONAL JOURNAL OF PLANT SCIENCES

the lower surface (fig. 5A) and narrow, coriaceous sun leaves (fig.
5B) with heavily papillate lower cuticles and deeply sunken sto-
mata. They were distinguished as two forms of the same species
(Kvaček in Kvaček et al. 2004, p. 29). Leaf morphology and epi-
dermal anatomy (papillate abaxial surface) both suggest a relation-
ship to the living Nyssa javanica complex.
The co-occurring fruits are preserved partly with the fleshy
exocarp (fig. 5E) and partly as isolated stones. The fruits in the
fleshy exocarps are similar to those of the Nyssa sylvatica–Nyssa
sinensis complex but are much larger. The endocarps are ribbed,
unlike those in N. javanica. According to Eyde (1997), they do
not match any living species.

Fig. 5 Nyssa bilinica (Ung.) Kvaček. A, Shade leaf (forma bilinica), Bı́lina Mine, northern Bohemia, Miocene, 31. B, Sun leaf (forma haidingeri),
same locality as A, 31.8. C, Abaxial cuticle of a shade leaf, same locality as A, 3300. D, Pollen in situ from staminate inflorescence, 31000. E, Fruit
with compressed exocarp, same locality as A, 31.8. F, Staminate inflorescence, same locality as A, 31.2.
 KVAČEK—WHOLE-PLANT RECONSTRUCTIONS OF FOSSIL ANGIOSPERMS 925

Staminate inflorescences (fig. 5F) with pollen in situ (fig. 5D)
co-occur in the same layer of the Bı́lina Mine and obviously be-
long to the same plant (J. Sakala, Z. Kvaček, and J. Dašková, un-
published manuscript). The inflorescences are globular (as opposed
to cylindrical in the N. sylvatica–N. sinensis group) and sub-
tended by small bracts, as in the N. javanica complex, but a
detailed comparative study of pollen in situ (Dasková 2000)
suggests a relationship to the N. sylvatica–N. sinensis group. In
this case, the fossil appears to represent an extinct Nyssa line-
age, perhaps endemic to the Miocene of Europe, that incorpo-
rates features of several living descendants from North America
and Asia. It is certainly not an isolated case in the European
Tertiary. Similar combinations of characters have been found
in extinct Juglandaceae (e.g., Engelhardia complex) and other
groups with the Asa Gray disjunction.
Cucurbitaceae
This last case, drawn from the Cucurbitaceae, shows how
combining co-occurring fossil organs may help guide a search
for living counterparts. The nicely preserved Pliocene flora of
Alsace collected by the late Fritz Geissert (Geissert 1972; Geis-
sert et al. 1990) is available as isolated carbonized leaf fossils.
Geissert wisely embedded the fossil leaf compressions in glyc-
erol on the bottom of transparent plastic boxes to keep them
moist for future cuticular studies (Kvaček et al. 2008).
One of the leaf forms (fig. 6A, 6C) is similar to Acer leaves
but is different in stomatal type and pubescence (fig. 6C). The
list of the co-occurring carpoflora (Geissert 1972) and prelimi-
nary reports (Geissert 1987; Geissert et al. 1990) include seeds
(fig. 6D) assigned to Trichosanthes fragilis E. M. Reid (Cucurbi-
taceae). This genus of herbaceous plants, which is today confined
to Asia and Australia, often produces palmately lobed leaves
somewhat similar to maple foliage. Comparison of epidermal
structure in Cucurbitaceae also shows that several leaf fossils
from Alsace, although quite variable in form, can be assigned to
genera of Cucurbitaceae near Trichosanthes on the basis of sto-
mata, trichomes, and peculiar shield glands on the leaf surface.
It is still uncertain whether the dispersed seeds really belong
to Trichosanthes or to a different genus of Cucurbitaceae. Most
authors (Szafer 1947; Dorofeev 1963, 1966; Geissert et al.
1990) followed the initial suggestion by Reid (1921), but per-
haps more comparative studies will be necessary to confirm this
systematic assignment. The seeds are characteristically orna-
mented on the surface but so far have not been compared ade-
quately because of the limited number of living plants. In leaf
architecture, the fossils compare well with only a few East

Fig. 6 Trichosanthes sp./Trichosanthes fragilis E.M. Reid. A, Isolated leaf compression, Auenheim, Alsace, Pliocene, 32.5. B, Isolated leaf
compression, Auenheim, Alsace, Pliocene, 32.2. C, Abaxial cuticle of the leaf in C, 3200. D, Seed, Sessenheim, Alsace, Pliocene, 33.
926 INTERNATIONAL JOURNAL OF PLANT SCIENCES
Asiatic living representatives of Trichosanthes that have entire-
margin–lobed leaves. Additional work on foliage of other ex-
tant Cucurbitaceae is needed to establish a more secure base for
further comparisons.
Mädler (1939) described leaf forms from the Pliocene of Nie-
derrad similar to those from Alsace as maple leaves (Mädler
1939, pl. 9, figs. 17, 18); these are accompanied by the kind of
seeds just described (Engelhardt and Kinkelin 1908; Mädler
1939). Unfortunately, this plant was completely destroyed by
bombing during the Second World War. Because of this newly
suggested reconstruction (Kvaček et al. 2008), Trichosanthes fo-
liage reported from the Upper Miocene of Kodor (Kolakovsky
1964) may not belong to this family. Those leaves correspond
much better to a different leaf form of the Miocene Berberidaceae
called Epimedium praeasperum (Andreánszky) Givulescu (1979).
Conclusions
Recent studies of fossil angiosperms reiterate the importance of
the whole-plant concept for a full understanding of fossil angio-
sperms. Although this can be a risky undertaking, reliable recon-
structions can be accomplished by the use several lines of evidence
(studies of pollen and cuticles in situ, co-occurring wood, foliage,
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can help correct errors in the systematic interpretation of previ-
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Acknowledgments
This article reviews results of many previous studies that I carried
out together with C. Bůžek, F. Holý, S. R. Manchester, H. Walther,
J. Sakala, and others. This fruitful cooperation is greatly appreci-
ated. I also greatly appreciate the opportunity to contribute to the
Else Marie Friis Jubilee Conference, Stockholm, 2007. I thank P.
Endress (Zurich) for discussion on relationship of Cercidiphylla-
ceae; J. Sakala (Prague) for data on fossil wood; S. R. Manchester
(Gainesville, FL) and the reviewers, who suggested improvements
of the text; and V. Teodoridis (Prague) for technical assistance in
preparing illustrations. Most work presented here was subsidized
by the Grant Agency of the Czech Republic within various projects,
most recently by the Ministry of Education and Youth, Czech Re-
public, within Research Scheme MSM 002162085.
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