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News from an old wood —Agathoxylon keuperianum (Unger) nov. comb.
in the Keuper of Poland and France
, Maria Barbacka
, Karoly Boka
, Pawel Filipiak
, Frédéric Thévenard
UniversitéLyon 1 and CNRS UMR 5276, 7 rue Dubois, F69622 Villeurbanne, France
Jagiellonian University, Institute of Botany, Department of Palaeobotany and Palaeoherbarium, ul. Lubicz 46, 31-512 Kraków, Poland
Faculty of Earth Science, University of Silesia, Będzińska 60 St, 41-200 Sosnowiec, Poland
Hungarian Natural History Museum, Department of Botany, 1476 Budapest,pf. 222, Hungary
W. Szafer Institute of Botany, Polish Academy of Sciences, Lubicz 46, 31-512 Kraków, Poland
Department of Plant Anatomy, Eötvös Loránd University, H-1117 Budapest, Hungary
Senckenberg Forschungsinstitut und Naturmuseum Frankfurt, Senckenberganlage 25, 60325 Frankfurt am Main, Germany
Senckenberg Center for Human Evolution and Palaeoenvironment, Institut für Geowissenschaften, Universität Tübingen, 72076 Tübingen, Germany
Received 8 April 2015
Received in revised form 10 June 2015
Accepted 11 June 2015
Available online 19 June2015
New fossil woods from the Keuper of Poland and France are assigned to Agathoxylon keuperianum (Unger) nov.
comb. A complete nomenclatural treatment of this taxon is given, with up-dated nomenclatural synonymy and
some taxonomical notes. It is shown that Zimmermann's choice of a lectotype (1953) for this species is
superseded by material in the Unger collection at the Paris Muséum National d'Histoire Naturelle (France).
Taxomical reappraisal of Unger collection also evidenced that Simplioxylon hungaricum Andreanzsky correct
name is Simplicioxylon wurtembergicum (Unger) nov. comb. and that Peuce brauneana Unger can be excluded
from Agathoxylon keuperianum synonymy. Literature data for Agathoxylon keuperianum are reviewed and some
are invalidated. Our new data are the ﬁrst safe reports for this species outside from Germany. The systematic
position of Agathoxylon keuperianum is discussed, as well as its stratigraphical and geographical range. In the
present state of knowledge it seems to be restricted to the areawhere German Keuper is deposited. Anatomical
featuressuggest that Agathoxylon keuperianum thrived under warmand wet conditions, whereas GermanKeuper
sediments globally suggest hot and dry climate.
© 2015 Elsevier B.V. All rights reserved.
After the seminal studies in the United Kingdom (Witham of
Lartington, 1831, 1833; Lindley and Hutton, 1832; Nicol, 1834),
Germany was the second cradle to the study of fossil wood, in partic-
ular with the important contributions of Göppert (1840, 1841, 1848,
1850, 1881),Unger (1847) and Hartig (1848).Amongtheﬁrst stud-
ied fossil woods, and hence the “oldest”ones, is the well-known
“Pinites keuperianus”(Unger, 1842; Braun, 1859). This wood is of in-
terest at least in three respects: 1) it has never been conﬁdently doc-
umented outside from Germany up to now, which is surprising for a
wood with the most ordinary Araucarioxylon-type of anatomy; 2) it
is known only from the Late Triassic, again a surprising ﬁgure for
such a type of wood; and 3) being bound to a sedimentary facies
known as the German Keuper, it potentially has a palaeoecological
Interestingly, as the anatomy of this taxon is quite characteristic,
many people used this wood to establish a Keuper age for the siliciﬁed
woods occasionally found redepositioned within Quaternary ﬂuvial
terraces in SW-Germany (Mägdefrau, 1960; Tyroff, 1978; Selmeier,
1994; Kelber, 2007).
Here we report Agathoxylon keuperianum (Unger) nov. comb. wood
from Upper Silesia, Poland, and Lorraine, France (Fig. 1). We provide a
full systematic treatment for this taxon. A new combination is proposed,
and its type is discussed. Eventually we discuss its systematic position
and its palaeobiogeography.
2. Geological and palaeoxylological setting
During the Triassic the Germanic Basin was a sedimentary basin
centered on Germany, stretching from Yorkshire (UK) to Southern
Poland (Fig. 2). Polish Triassic sedimentary basin is part of the Germanic
Basin (Mader, 1997; Franz, 2009), however its marginal position, and
Review of Palaeobotany and Palynology 221 (2015) 83–91
⁎Corresponding author at: 7 rue Dubois, F69622 Villeurbanne cedex.
E-mail addresses: firstname.lastname@example.org (M. Philippe), email@example.com
(G. Pacyna), firstname.lastname@example.org (Z. Wawrzyniak), email@example.com
(M. Barbacka), firstname.lastname@example.org (K. Boka), pawel.ﬁlipiak@us.edu.pl (P. Filipiak),
email@example.com (L. Marynowski), firstname.lastname@example.org (F. Thévenard),
Dieter.Uhl@senckenberg.de (D. Uhl).
0034-6667/© 2015 Elsevier B.V. All rights reserved.
Contents lists available at ScienceDirect
Review of Palaeobotany and Palynology
journal homepage: www.elsevier.com/locate/revpalbo
marked local facies variability make stratigraphical correlations dif-
ﬁcult (Szulc et al., 2006; Racki, 2010; Pacyna, 2014). A new frame-
work was recently proposed by Szulc and Racki (2015).They
referred most Upper Silesia Keuper strata to the redeﬁned Grabowa
Formation. Al l Polish samples investigated here (fro mPatoka, Poręba
and Marciszów-Zawiercie) derive from the Marly Mudstone–Sand-
stone member of the Grabowa Formation (lateral equivalent to the
German Arnstadt Formation). They were found in variegated mud-
stones, sandstones and carbonates of Norian age. Based on palynologi-
cal data, wood source strata were referred to the local Polish Corollina
meyerana palynozone (Orłowska-Zwolińska, 1983, 1985) which is cor-
related with Middle Europe Granuloperculatipollis rudis zone (Cirilli,
2010; Kürschner and Herngreen, 2010). Investigated localities provided
rich invertebrate and vertebrate tracks and skeletal remains (Szulc
et al., 2006; Dzik et al., 2008; Budziszewska-Karwowska et al., 2010;
Racki, 2010; Skawina and Dzik, 2011; Sulej et al., 2012; Sadlok and
Polish Keuper fossil wood has not yet been taxonomically studied in
great detail. Fossil woods from Zawiercie area were already described
by Lilpop (1917) as “Blanowicer Keuperholz”, on the basis of specimens
from the “Elka”,“Kamilla”and “Zygmunt”coal pits. The specimens are
said to be xylologically similar, but Lilpop did not assign them to a
named taxon, his description is not accurate enough and illustration is
limited to a cross-section. Unfortunately Lilpop's collection has not
been preserved. Lower Jurassic (Upper Pliensbachian) strata crop out
in the Zawiercie area. Coal seams of low economic value were mined
in these strata during the 19th and early 20th centuries. According to
Różycki (1930) some thin Keuper coal seams have also been exploited
in Zawiercie area, it is not possible hence to rule out that Lilpop's spec-
imen derived from Keuper coal seams. Neverthelessthe levels mined in
the three mentioned pits are now recognized as Liassic in age (Pacyna,
Petriﬁed fossil wood from Poręba were reported by Brzyski and
Heﬂik (1994) and assigned to Araucarioxylon sp. They were found ex-
situ in a stream, and derive from the Woźniki limestones, which are
Norian in age. Their cross-ﬁeld pits are not preserved, however the
other xylological characters are similar to that of the here described
North-eastern France (Lorraine, Franche-Comté and part of the Bassin
de Paris) was also part of the Germanic Basin (Mader, 1997). It was locat-
ed less marginally as compared to Poland, however, and its sedimento-
logical succession is usually considered typical. French Keuper wood
has been little studied. Fliche (1910) reported Agathoxylon keuperianum
from the Keuper of Lorraine, his descriptions and illustrations being in-
accurate, and sample age being the main identiﬁcation argument. The
Xenoxylon conchylianum wood from Vaubexy (Vosges, NE France) origi-
nally indicated as Muschelkalk (Middle Triassic) in age by Fliche (1910)
is actually originating from the Carnian (Philippe and Thévenard,
1996). On the meager basis of these two data, French Keuper fossil
wood record is less diverse but otherwise similar to the German one
3. Material and methods
The new material originates fromPoland (Patoka, Poręba, Zawiercie-
Marciszów) and France (Provenchères, Varangéville) (Fig. 1). The
French material was discovered in existing palaeontological collections.
3.1. Upper Silesia sites
Woody axes are numerous in Patoka locality, mostly preserved as
coaliﬁed(jet-like) fossil wood and less commonly occurringas charcoal
fragments. Their dimensions vary in length from several cm to one
meter or more. They are derived from the greenish/yellowish some-
times gray layers of mudstone/siltstone with sandstone lenses of the
Fig. 1. Localities for newly reported Agathoxylon keuperianum nov. comb. material, in France (A) and in Poland (B).
Fig. 2. Distribution mapfor Agathoxylon keuperianum, ona sedimentological background.
In situ occurrences marked by stars. Map based on Ziegler (1990).
84 M. Philippe et al. / Review of Palaeobotany and Palynology 221 (2015) 83–91
middle portion of the Patoka Member which crops out in the Patoka
Brickyard pit (see Szulc and Racki, 2015). Well-preserved coniferous
leafy shoots were also found associated with stems. Several charcoals
from this locality are kept at Jagiellonian University, Institute of Botany,
Department of Palaeobotany and Palaeoherbarium, among which KRA-
PALEO 104/200 was used to prepare illustration.
At this locality charcoal is quite abundant and co-occurs with
coaliﬁed and calciﬁed fossil wood. Charcoal fragments are usually
large (up to 5 cm of diameter), very well preserved due to early-
diagenetic mineralization and low compaction of parent sediments
(Marynowski et al., 2014; Kubik et al., in press a,b). So far a number of
charred wood with anatomically different morphotypes were found
here (Kubik et al., in press a,b). Based on palynomorphs this locality
can be dated as being of mid–late Norian age (Sulej et al., 2012;
Niedźwiedzki et al., 2014). Poręba locality revealed a rich Upper Triassic
faunal and ﬂoral assemblage containing e.g., turtles, dinosaurs, sharks,
conifers shoots and probable ginkgoalean leaves (Sulej et al., 2012;
Niedźwiedzki et al., 2014).
Charcoals and fossil wood remains preserved as calciﬁed or coaliﬁed
fragments are common at this site. This locality yielded several charcoal
specimens which can be assigned to Agathoxylon keuperianum.Basedon
the composition of the vertebrate fauna, which includes large dicyno-
donts and archosaurs (Budziszewska-Karwowska et al., 2010), this
locality can be correlated with other localities from the same region
dated as uppermost Norian–lower Rhaetian (Dzik et al., 2008;
Niedźwiedzki et al., 2012). In addition, fresh water bivalves, teeth
of sarcopterygian ﬁshes and remains of horsetails and conifers have
been found there (Szulc et al., 2006; Budziszewska-Karwowska
et al., 2010; Racki, 2010; Skawina and Dzik, 2011; Sadlok and
All specimens from Zawiercie-Marciszów are ex situ, and were col-
lected from a dump of Upper Triassic strata, excavated during the con-
struction of the city landﬁll site (Sadlok and Wawrzyniak, 2013).
These specimens include WNOZ/S/7/247, WNOZ/S/7/248, WNOZ/S/7/
249 and WNOZ/S/7/250, all held at the University of Silesia Faculty of
3.2. French sites
Sample MP11 was found in the Palaeontological collections of the
Besançon University in 1988. Its label states that it originates from the
Upper Rhaetian of Provenchères (Haute-Marne, France). In this north-
eastern France Rhaetian locality, at the rim of the Bassin de Paris, typical
Germanic Keuper crops out. Grindstones were prepared there, in
quarries which exploited ﬁne-grained sandstone interbedded with var-
iegated clay. This fossil-rich localityis well-known as it yielded an abun-
dant fauna, including Prosauropod dinosaurs, plesiosaurs, ichthyosaurs,
sharks, and ﬁshes (Cuny, 1995). Plants, however, had never been re-
ported with accurate taxonomic assignment. Palaeoenvironment is
usually interpreted as high-energy shallow-marine.
A second set of three samples (n°3292/MP1830) was located in the
Joly's collections of the Ecole Nationale Supérieure de Géologie
(Nancy, France). It was collected by Henry Joly (1881–1975) who was
teaching geology at the Nancy University (Maubeuge, 1978). It origi-
nates from Varangéville (Meurthe-et-Moselle, France), from a similar
stratigraphical (“Grès bigarrés rhétiens”), sedimentological (sandstones
layers interbedded with variegated clays) and geographical (Bassin de
Paris rim) context (Joly, 1937). There also a rich palaeontological
content was recorded, including reptilian, therapsid and mammalian
teeth (Godefroit, 1997). Varangéville palaeoenvironment is usually
interpreted as more proximal than that of Provenchères, possibly
Beside the French and Polish material we also use here slides kept at
the Museum für Naturkunde–Leibnitz Institut für Evolution und
Biodiversität Berlin (Germany), under the numbers 360, 366 and 368.
This material, partly illustrated by Gothan (1908), originates from
Heldburg near Coburg (Germany), a classical locality for Keuper
palaeobotany, and is labeled “Dadoxylon keuperianum Göppert”.
Fossil wood was studied with several techniques, including Collodi-
on microcasts and SEM (Philippe et al., 2006). The illustrations present-
ed here were prepared in Poland at the Silesia University Faculty of
Earth Science, using an environmental scanning electron microscope
(ESEM) Philips (Plates I and II), and in Hungary at the Eötvös Loránd
University Department of Plant Anatomy, using standard SEM
techniques (gold coated samples) and a Hitachi-2360 N microscope
4.1. Systematic treatment
Genus Agathoxylon Hartig, Bot. Zeitung (Berlin) 6: 190. 10 Mar 1848.
Type: A. cordaianum Hartig.
Species Agathoxylon keuperianum (Unger) Philippe nov. comb.
1842 Pinites keuperianus Unger, Chloris Protagaea pars 2/3: 31 (diagno-
sis, no illustration).
Selected nomenclatural synonyms (complete list in Vogellehner, 1965):
1847 Dadoxylon keuperianum (Unger) Endlicher, Synopsis coniferarum:
1848 Araucarites keuperanus (Unger) Goeppert in Bronn, Gesch. d. Nat.
III 2: 91.
1870 Araucarioxylon keuperianum (Goeppert) Kraus in Schimper, Traité
de Paléontologie végétale: 384.
1908 Peuce keuperina Engel, Geognostischer Wegweiser durch
1911 Pagiophyllites keuperianus Tuzson, Monographie der fossilen
Pﬂanzenreste der Balatongegend: 31.
Nomenclatural notes: Different synonymies are available in the
literature, some of them are partly erroneous. Vogellehner (1965) ﬁrst
clariﬁed the situation, evidencing that the original publication was by
Franz von Unger (1842: 31) in the second issue (parts 2 and 3) of his
Chloris Protagaea, later published as a complete volume in 1847. Ac-
cording to Göppert (1848: 91) this name was ﬁrst proposed by himself
in a manuscript before 1842. It could thus be possible to use Agathoxylon
keuperianum (Göppert ex Unger) nov. comb. Although using “keuperanus”
in 1848 Göppert switched to “keuperianus”in 1850. Hartig knew about
Agathoxylon keuperianum (Hartig, 1848: 140) but assigned it to Dadoxylon
on the belief that the wood had no axial parenchyma. In 1908 Engel indi-
cated as a synonym “Peuce keuperina aut.”(sic), although no evidence was
found in the scientiﬁc literature of this name being used.
Locus et stratus typicus: Keuper sandstones in Attelsdorf, near Bamberg
(north of Nuremberg, Bavaria, Germany).
Typus: Three slidesfrom Göppert's Arboretum Fossile (a collection of fos-
sil wood slides which was sold by Göppert around 1881) were chosen
by Zimmermann (1953) as “lectotype”. There is no information, howev-
er, about the log from which these slides were prepared. Unger gave no
indications about the number of specimens he studied, neither in his
protologue (1842) nor in his further publications (1847, 1850).
Vogellehner (1965) concluded that further investigations should be
performed in order to locate the original material. In Paris, at the
Muséum National d'Histoire Naturelle (MNHN), the Unger collection
includes slides 8770, labeled “Attelsdorf, Pinites keuperianus”. This can
85M. Philippe et al. / Review of Palaeobotany and Palynology 221 (2015) 83–91
be interpreted either as the holotype or as the only syntype left. Its
xylology ﬁts well with what is named Agathoxylon keuperianum today.
Taxonomical synonymy: In his Chloris protagaea Unger (1842) also de-
scribed Peuce brauneana, mentioning material from the Upper Keuper
of Kulmbach, and from the early Jurassic near Bayreuth, Germany.
Vogellehner (1965: 15) assigned, with doubts, P. brauneana to
Agathoxylon keuperianum, simultaneously proposing Dadoxylon
brauneanum (Unger) Vogellehner as a nov. comb. Vogellehner, how-
ever, did not review the Unger collection. Within the Unger collection
at the MNHN three slide sets are labeled “Peuce Brauneana”:slides
Plate I. Specimen WNOZ/S/7/247. Fig. 1 cross-section, general view, homoxylous wood, growth-rings little marked. Fig. 2 cross-section,early-wood tracheids with quadrate to polygonal
section. Fig. 3 tangential view, dominance of low rays. Fig. 4 half-tangential longitudinal view, araucarian radial pit rows and homogenous rays with araucarioid cross-ﬁelds. Fig. 5 radial
view homogenous rays, marginal rays beinglocally larger. Fig. 6 radial view, cross-ﬁelds with 1 to 5 oculipores each, late wood on the left.
86 M. Philippe et al. / Review of Palaeobotany and Palynology 221 (2015) 83–91
228, 229, and230; slide 231; and slide232. Only the ﬁrst set is labeled as
originating from the “Oberer Keuper von Culmbach bei Bayreuth”. Slide
231 label indicates “Unterer Lias”and slide 232 hasno stratigraphical in-
formation. From our review of these three slide sets P. brauneana Unger
is heterotypic, slides 231 and 232 being a Brachyoxylon (sensu Philippe
and Bamford, 2008), whereas slides 228, 229 and 230, all three made
from the same specimen, are referable to Agathoxylon keuperianum.
We here lectotypiﬁed P. brauneana Unger with slide 232 in the Unger
collection at the MNHN, in order to avoid nomenclatural instability for
the well known Agathoxylon keuperianum. Nevertheless the material
assigned by Braun to P. brauneana is mostly referable to Agathoxylon
keuperianum (Kraus, 1866; Vogellehner, 1965).
Tuzson (1911) included Peuce wurtembergica Unger and Araucarites
thuringiacus Bornemann in Agathoxylon keuperianum. The syntypes for
P. wurtembergica in the Unger collection at MNHN (slides 166–171)
are variably preserved. Only slides 166 and 167 are well enough
preserved for xylological investigations. They display thesame features
as Simplicioxylon hungaricum Andreánszky (1949): 250, text-ﬁg. 1–6,
Toarcian, Urkut, Hungary, the correct name of which is hence
Simplicioxylon wurtembergicum (Unger) nov. comb. Vogellehner
(1965) kept A. thuringiacus Bornemann as an Agathoxylon keuperianum
taxonomical synonym, although he did not review Bornemann's type.
We neither were able to access this material.
Roselt (1986: 696) suggested Araucarioxylon mineense Ogura as
a taxonomical synonym of Agathoxylon keuperianum. This Triassic
Japanese wood is however a Protocedroxylon (Nishida and Oishi, 1982).
4.2. Description of the new material and identiﬁcation
All studied samples have the same anatomy, although anatomically
variable and variously preserved. The material WNOZ/S/7/247 from
Zawiercie Marciszów is described and illustrated ﬁrst (Plate I): the
wood is a homoxylous tracheidoxyl (Plate I, Fig. 1); some narrower tra-
cheids suggest growth-rings with narrow late wood (2–3cells wide);
early wood tracheids are mostly quadrate in cross-section and thin-
walled (Plate I, Fig. 2); rays are 1–11 cells high, mostly 3–5 cells high
(Plate I, Fig. 3, 4), homogenous, marginal cells being however, some-
times larger (Plate I, Fig. 5); radial pitting is uniseriate, in long araucar-
ian chains (Plate I, Fig. 4); these pits are contiguous (appearing
sometimes distant depending on preservation type), round or slightly
ﬂattened; radial pit pori are somewhat enlarged by charring; ray cell
walls are thin and, except for cross-ﬁelds, unpitted (Plate I, Fig. 5, 6);
cross-ﬁelds are araucarioid, with 1–4 (9) large alternate contiguous
cupressoid oculipores (Plate I, Fig. 6), quite often with a single large
pit; and resiniferous axial parenchyma occurs locally, however no
resin canal was observed.
Plate II. Specimen WNOZ/S/7/248. Fig. 1 cross-section, thin late-wood, regular tracheid pattern. Fig. 2 cross-section, thin-walled early wood tracheids with irregular pattern. Specimen
WNOZ/S/7/249. Fig. 3 tangential view, inﬂated ray, probably charred when still water containing. Fig.4 araucarian radial pit rows modiﬁed by charring.
87M. Philippe et al. / Review of Palaeobotany and Palynology 221 (2015) 83–91
The rest of the material ﬁts with this description and allows a more
comprehensive view of the taxon. Late wood is thin, limited to a few tan-
gentially ﬂattened cell layers (Plate II, Fig. 1). Early cell walls are strikingly
thin, tracheid cross-section pattern being locally irregular (Plate II,Fig.2).
In some of the charcoals inﬂated rays are often observed (Plate II,Fig.3).
Such happens when relatively fresh and wet wood burns at low temper-
ature (e.g., Rossen and Olson, 1985; Schweingruber, 2001). Charring en-
larged pit pori, but also made fainter their areolae, although well
preserved pits are always clearly araucarian (Plate II, Fig. 4). Over 90%
of observed cross-ﬁelds are of the araucarioid type described above,
with 2–6 cupressoid alternating contiguous oculipores. Locally, however,
cross-ﬁeld pitting is strikingly variable (Plate III, Fig. 1). Mostly in
narrowest tracheids, but not only, cross-ﬁelds sometimes have a single
oculipore (Plate III, Fig. 2). Locally this pit gets radially stretched, mimick-
ing a Xenoxylon window-like pit, but remaining areolated (Plate III,Fig.1,
3). In some cross-ﬁeld oculipores are not homogenous (Plate III,Fig.4).
This cross-ﬁeld variability emphasizes that by any fossil softwood a
large number of cross-ﬁelds must be studied to encompass natural
As for identiﬁcation, with araucarian radial pitting, araucarioid
cross-ﬁelds and thin and unpitted ray-cell walls these woods belong
to Agathoxylon Hartig (Philippe and Bamford, 2008). Within this genus
they ﬁt well within Agathoxylon keuperianum, a taxon known since
long and well illustrated by Vogellehner (1965:Plate I Fig. 9 and Plate
II Figs. 10–18). The peculiar feature illustrated by Vogellehner (op. cit.
Plate II Fig. 14), with 4 oculipores in a radial row, is very uncommon
by araucarioid cross-ﬁelds, and was also observed by the Polish material
(sample KRA-PALEO 104/200). Although typical araucarioid cross-ﬁelds
aremostfrequentbyAgathoxylon keuperianum, this taxon is well-
known for its signiﬁcant crossﬁeld percentage with a single large
oculipore (Mägdefrau, 1960; Vogellehner, 1965), which is alsoobserved
here. Cross-ﬁeld pitting being remarkably variable by this wood, limited
observation could lead to wrong identiﬁcation. This variability also sug-
gests that several Agathoxylon taxa recognized from the German Keuper
might represent atypical or superﬁcially observed specimens of
In Berlin we chanced upon two Agathoxylon keuperianum slides with
preserved pith (n°366 and 368). By both specimens pith reaches 1.5 cm
in diameter (1.4 and 1.7 cm respectively), and include more than a hun-
dred primary xylem strands. By both specimens pith issurrounded by a
secondary xylem cylinder (6.7 and 6.6 cm in diameter respectively),
with concentric andaxially symmetric growth rings, suggesting vertical
axes. Albeit such large piths are not unknown by modern conifers they
are rare and can be considered as typical for growth under optimal
Original data for Agathoxylon keuperianum were listed by
Vogellehner (1965: 15–16).Table 1 sums up some data not
Plate III. Specimen KRA-PALEO 104/200, Jagiellonian University, Institute of Botany, Department of Palaeobotany and Palaeoherbarium. Fig. 1 radial view, cross-ﬁeld variability (cross-
ﬁelds preserved as calcite ﬁllings or on original tracheids). Fig. 2 radialview, single oculipored late wood cross-ﬁelds. Fig. 3 radial view, Xenoxylon-like elongated oculipores. Fig. 4, radial
view, cross-ﬁeld with 3 oculipores, on of then signiﬁcantly larger.
88 M. Philippe et al. / Review of Palaeobotany and Palynology 221 (2015) 83–91
mentioned in Vogellehner's list. The Late Triassic fossil wood
record of Europe is limited, with a large majority of data from the
Keuper of Germany (Zimmermann, 1953; Vogellehner, 1965,
1967). Zimmermann (1953) described three taxa (Dadoxylon
gaildorﬁanum,Dadoxylon graminovillae and Dadoxylon implexum)
co-occurring with Agathoxylon keuperianum in the Stubensandstein
Württemberg of SW Germany (Upper Keuper, Norian age). From
their protologue these four taxa are xylologically similar, and should
they be reuniﬁed, and would be named Agathoxylon keuperianum.
Vogellehner (1965) evidenced the occurrence of diverse Keuper
wood ﬂoras from Southern Germany (Franconia and Southern Thu-
ringia), including e.g., Xenoxylon Gothan and Protocupressinoxylon
Eckhold, beside Agathoxylon.
There is a report of Agathoxylon keuperianum from north-eastern
France (Vic-sur-Seille, Moselle; Fliche, 1910, see Table 1). Vogellehner
(1965) suggested assigning Fliche's material to a new taxon, he
named Dadoxylon alsaticum. This name is not validly published, howev-
er, as Vogellehner included in the taxon two sets of material without in-
dicating a type (ICBN, art. 40.1, Mcneill et al., 2012). Fliche's description
and illustration are not detailed enough to substantiate beyond doubt of
his identiﬁcation. Despite careful researches in Nancy, were Fliche
taught, and elsewhere in France we never succeed in locating Fliche's
collection. At this stage this data must be considered as doubtful.
McLean (1926) described a Mesembrioxylon rhaeticum from the
Rhaetian of Wales, interpreting altered araucarioid cross-ﬁelds as
“obliquely elongated simple pits”. Should the other features having
been correctly interpreted, M. rhaeticum probably belongs to
Brachyoxylon Hollick & Jeffrey. It originates from the Penarth Group, a
sedimentary set of gray, marine mudstones and limestones, dramatical-
ly contrasting withthe underlying levels, typical for German Keuper fa-
cies. Also from Wales, Protocupressinoxylon rhaeticum Vogellehner,
based on Cheirolepis associated charcoal from supposedly Rhaetian ﬁs-
sure ﬁllings (Harris, 1957), is probably conspeciﬁc with M. rhaeticum.
Mathiesen's collection in Copenhagen (Denmark) includes a lot of fossil
woods from the Late Triassic of Jameson Land, Greenland (work in prog-
ress), however none of these ﬁtw
According to Greguss (1967) a wood from the Permian of
Hungary is similar to Agathoxylon keuperianum,butthematerial
kept in the Hungarian Geological Survey does not support, however,
To sum up the Late Triassic wood record in Western Europe is largely
concentrated in Germany. There, Agathoxylon keuperianum is dominant,
together with some closely similar taxa (D. gaildorﬁanum, etc.),
associated with much rarer wood types (Protocupressinoxylon,
Xenoxylon). Agathoxylon keuperianum had not yet been conﬁdently doc-
umented outside of the German Keuper distribution area.
5.1. Systematic relationships
According to Greguss (1967) the ex-situ Agathoxylon keuperianum
described by Mägdefrau (1960) could be assigned to Baieroxylon,a
genus he explicitly related to Ginkgoales. The features Greguss consid-
ered as diagnostic for Baieroxylon are, however, found in various
Palaeozoic and Mesozoic woods with no afﬁnities to the Ginkgoales.
Schimper (1872) hypothesized, on the basis of common association,
that Agathoxylon keuperianum is the wood of Glyptolepis,whereas
Kraus (1866) and Hoffman (1884) underlined that it is often associated
to abundant Voltzia coburgensis, probably the most characteristic plant
remain of the German Keuper Mägdefrau (1953).Tuzson (1911)
assigned the wood to a new genus, Pagiophyllites, considering safe that
it is the wood of a Pagiophyllum. More recently Voltzia was proposed
to be the plant genus that yielded this Agathoxylon keuperianum wood
Within the German Keuper Agathoxylon keuperianum is often found
associated to conifer foliages, especially Voltziales, however occasional-
ly also with other gymnosperm groups (Kelber, 2007).
From its anatomy it is impossible to unambiguously relate Agathoxylon
keuperianum to any modern conifer group. Despite the obvious etymolo-
gy, assignment of a fossil wood to Agathoxylon does not imply per se that
it was the wood of an Araucariaceae. Some features observed here, and
especially the high number of cross-ﬁelds featuring a single oculipore
(which is sometimes much elongated) are unknown by the extant
Araucariaceae. Moreover the hypothesis is quite unrealistic that modern
conifer families had a xylological diversity which remained unchanged
all through the Mesozoic and Cenozoic. Palaeochemotaxonomy could
give some hints (Otto and Wilde, 2001; Marynowski et al., 2007,2008),
but only calciﬁed fossil wood seems to be perspective.
The variability of Agathoxylon keuperianum cross-ﬁeld pitting is strik-
ing. The local occurrence of Xenoxylon-like elongated cross-ﬁeld pits is
of interest. Indeed, it suggests that the conifer group which built
Agathoxylon keuperianum wood had high xylological diversity, and thus
might have been basal to several modern conifer families. Another
wood with variable cross-ﬁeld pitting is Protocupressinoxylon purbeckensis
Some Agathoxylon keuperianum (Unger) nov. comb.data not listed in Vogellehner (1965). Not all originalmaterials were controlled and thus some of these datamight be doubtful. Fliche's
data e.g.,is poorly substantiatedand type material wasnot reappraised. Numerous mentions on the WWW are notbased on xylologicalinvestigations andthus are not taken into account
Reference Stratigraphy Locality
Hoffman (1884) Keuper Coburg area, Bavaria, Germany
Vater (1884) Ex-situ, Oligocene (?) or
Braunschweig, Lower Saxony, Germany
(possibly same site as for Gottwald, 1992)
Schuster (1909) Carnian Neustadt an der Aisch-Herrnneußes, Bavaria, Germany
Fliche (1910) “Marnes irisées inférieures”, Lower Keuper Vic-sur-Seille, Moselle, France
Linck, 1949; Kelber, 2007; Mohr et al., 2008 Middle Keuper Streufdorf, Thuringia, Germany
Mägdefrau (1960) Ex situ. Quaternary sediments Bavaria, Germany
Tyroff (1978) Ex-situ Hesse, Germany
Gottwald (1992) Ex-situ Lower Saxony, Germany
Selmeier (1994) Ex-situ Bavaria, Germany
Kelber (2007) Rhaetian Heigelsdorf, Bavaria, Germany
Kubik et al., in press a,b Norian Zawierce-Marciszów, Upper Silesia, Poland
This study Upper Rhaetian Provenchères, Haute-Marne, France
This study Rhaetian Varangéville, Meurthe-et-Moselle, France
This study Norian Patoka, Upper Silesia, Poland
This study Norian Poręba, Upper Silesia, Poland
This study Norian Zawiercie-Marciszów, Upper Silesia, Poland
89M. Philippe et al. / Review of Palaeobotany and Palynology 221 (2015) 83–91
Francis, from the Purbeckian of Western Europe (Philippe et al., 2010). It
grew similarly in lowlands with highly tropophilous sub-tropical climate.
5.2. Agathoxylon keuperianum distribution
From Table 1 and Vogellehner (1965) Agathoxylon keuperianum is
distributed stratigraphically only within the Keuper (Late Triassic), i.e.,
in the Carnian, Norian and Rhaetian stages. It has not yet been reported
from the Muschelkalk (Middle Triassic), nor from any Early Jurassic de-
posits. In Europe, the Muschelkalk fossil wood record is limited
(Vogellehner, 1967; Grauvogel-Stamm et al., 2003), and the Hettangian
one just somewhat richer (Philippe, 1995; Philippe et al., 2006)which
limits the inference that Agathoxylon keuperianum could be used as a
Keuper index-fossil, including dating ex-situ reworked siliciﬁed wood.
The wood geographical distribution also seems limited (Fig. 2). Late
Triassic wood record is, again, limited in several countries. Nevertheless,
Agathoxylon keuperianum is not reported from the Rhaetian of Green-
land where S. wurtembergicum is abundant (work in progress). It is
also absent from the Late Triassic fossil wood ﬂora of the United
Kingdom, northwestern and southern France. In the present state of
knowledge Agathoxylon keuperianum seems limited to the German
Keuper Basin, a geological unit recognized since long on a sedimento-
The wood Agathoxylon keuperianum is found within a wide range of
sediment types: ﬂuvial and marine sandstones, marine tempestites
(Provenchères), and variegated clays. As it was never found rooted in-
situ, its original ecology is not known. Being largely sedimentologically
distributed, from terrestrial to marine settings, it can be hypothesized
to have grown alongside coastal streams rather than really inland,
where its mobilization would have been difﬁcult.
During the Keuper, Europe was located at about 15°N, and experi-
enced a tropophilous tropical climate. This is usually considered as con-
tinental and arid, although it is thought to have evolved wetter by the
end of the Keuper Keuper (Bonis and Kürschner, 2012). Palaeogeo-
graphically the Keuper sediments deposited in a wide ﬂat area. These
were mostly siliciclastic, brought into the sea by large ﬂat deltas and
widely redistributed by sea currents and winds. Flash ﬂoods might
have occurred, as during the Carnian Pluvial Event (Simms and Ruffel,
1989), succeeded by times of intense evaporation with halite deposi-
tion. Although clearly peculiar from a sedimentological point of view
The German Keuper Basin does not seem to have a peculiar ﬂora
(Dobruskina, 1994), however a detailedsynecological approach still re-
mains to be performed.
Climate probably explains both the sedimentological peculiarities of
the German Keuper Basin and why Agathoxylon keuperianum distribu-
tion was limited to this basin. The wood features, however, cannot un-
equivocally be interpreted for climatological inferences. Growth-ring
are poorly marked, with a faint late-wood, however this is a common
feature by several Mesozoic and modern conifers, with no particular cli-
matological meaning (Brison et al., 2001). The tracheids are neither par-
ticularly broad nor narrow; nevertheless at leastsome of the early wood
tracheid walls are unusually thin, suggesting quick growth and unlimit-
ed water availability. The high frequency of single-pitted cross-ﬁelds
might be interpreted in climatological terms. Indeed, cross-ﬁelds with
numerous small oculipores might be hypothesized as differing in their
ability to propagate embolisms as compared to single-pitted cross-
ﬁelds. Currently available literature, however, does not give clear indi-
cations on this subject, however, and moreover the different types of
cross-ﬁelds seem to be distributed stochastically in the wood. Eventual-
ly the observations made on the Berlin specimens pith are probably giv-
ing the safest clue for a growth under much favorable circumstances,
from both a temperature and water availability point of view.
Of note, at generic level France and Germany have similar Keuper
wood ﬂora, with Agathoxylon and Xenoxylon.Vogellehner (1965) also
reported Protocupressinoxylon sp. from the Keuper of Franconia, but
corresponding wood is poorly preserved and protologue illustration
suggests Steinkern preservation bias. Xenoxylon is known in the Middle
Jurassic of Poland (Philippe et al., 2006) but has not yet been docu-
mented from the Triassic rocks there. The record is too spotty to inter-
pret this difference. Xenoxylon record starts within the Late Triassic, in
a palaeolatitudinal belt stretching from Europe to Japan (Oh et al.,
2015). Whereas eastward of this distribution during the Late Triassic
Xenoxylon is associated to Protocedroxylon Gothan, in the German
Keuper Basin it is associated with Agathoxylon keuperianum. Again this
might point out some climatological peculiarity of the German Keuper
Basin at that time.
Eventually, both the Xenoxylon and Agathoxylon keuperianum oldest
records are from the Carnian Schilfsandstein (Schuster, 1909;
Vogellehner, 1965), a deposit that represents the Carnian Pluvial
Event. Xenoxylon being indicative of cooler/wetter climate (Oh et al.,
2015), it is possible that Agathoxylon keuperianum required signiﬁcant
rain amounts, which it did not ﬁnd outside the German Keuper Basin.
Gilles Cuny kindly shared his knowledge of north-eastern France
Upper Triassic. Fabrice Malartre's help while searching the Nancy
palaeontological collection is much appreciated, as well as his informa-
tion about the Carnian Pluvial Event. Dario de Franceschi help in Musé-
um National d'Histoire Naturelle was much appreciated, just as that of
Daniel Contini in Université de Besançon palaeontological collections.
This work was partly supported by funds from the Polish National Sci-
ence Centre (grant No DEC-2012/05/B/NZ8/00990) and statutory
funds of the W. Szafer Institute of Botany, Polish Academy of Sciences.
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