Fungusplant interaction in a Thuringian (Late Permian)

Dadoxylon sp. in the SE Iberian Ranges, eastern Spain

Carmen Dieguez
a,
*
, Jose Lo pez-Go mez
b
a
Museo Nacional de Ciencias Naturales, CSIC, Jose Gutierrez Abascal, 2. 28006 Madrid, Spain
b
Instituto de Geolog a Econo mica-Dpto. Estratigraf a, CSIC-UCM, Facultad de Geolog a, Universidad Complutense, 28040 Madrid, Spain
Received 9 January 2004; received in revised form 16 November 2004; accepted 24 June 2005
Abstract
This report describes an anatomical and taphonomical study of a large, exceptionally well-preserved Late Permian Dadoxylon
sp. trunk. The specimen, found in siliciclastic red bed sediments of fluvial origin of the Landete Formation, Iberian Ranges,
eastern Spain, includes the first described fungusplant interaction for this time interval in Europe. This finding is all the more
significant since a bfungi-enriched layerQ, recently reported in sediments of different continents corresponding to the end of the
Permian, has been related to a world-wide crisis, whereby fungi were responsible for the breakdown of massive amounts of
vegetationalready suffered by a possible catastrophic event or eventsleading to a global drop in photosynthesis. The well-
known humid, temperate climatic conditions of the Late Permian sediments and the sedimentary environment represented by the
Landete Formation would have favoured the proliferation of fungi which was probably unaffected by any other significant biotic
activity. Fungal activity is likely to have started soon after the fall of the plants, indicated by the good preservation state of areas of
the trunk unaffected by the fungal infection. Later diagenesis suggests different fluid migration stages, clearly conditioned by the
original anatomical characteristics of the trunk and the results of fungal activity. Physical signs left by the enzyme action of wood-
decay fungi can be recognised in the fossilised wood. Using this information and sedimentological data, we reconstructed the
taphonomic processes incurred by the specimen, including both biostratinomic and diagenetic stages.
Ullmania sp. and pollen grains such as Lueckisporites and Nuscoisporites dulhuntyi produced by species of Late Permian
Walchiaceae or Ullmanniaceae that did not cross the Permian-Triassic boundary were also found in levels stratigraphically
below the fossil specimen; none occurred above the trunk site. This flora and fungal activity could relate this level of Iberian
Permian sediments to the level or one of the levels associated with the worldwide Late Permian crisis and fungal proliferation.
D 2005 Published by Elsevier B.V.
Keywords: Permian-Triassic boundary; Iberian Ranges; Wood-decay fungi; Dadoxylon
1. Introduction
Recent advances in research into the Permian-
Triassic include the demonstration that different
0031-0182/$ - see front matter D 2005 Published by Elsevier B.V.
doi:10.1016/j.palaeo.2005.06.031
* Corresponding author.
E-mail address: mcncd722@mncn.csic.es (C. Dieguez).
Palaeogeography, Palaeoclimatology, Palaeoecology 229 (2005) 6982
www.elsevier.com/locate/palaeo
events of considerable palaeoenvironmental signifi-
cance might be related to the end of Permian or to
the Permian-Triassic transition. These events mainly
involved changes in atmospheric characteristics,
marked volcanic events, superanoxia in oceans, and
mass marine and continental extinctions (see Isozaki,
1994; Retallack, 1995; Hallam and Wignall, 1997;
Kozur, 1998; Yin and Tong, 1998; Looy et al.,
Fig. 1. Upper part: geographical and geological location of the study area. Numbers indicate the main present-day basin, ranges and the Iberian
Massif. Lower part: palaeogeographical sketch of the western Tethys area during the Late Permian. Modified from Ziegler (1988).
C. Dieguez, J. Lo pez-Gomez / Palaeogeography, Palaeoclimatology, Palaeoecology 229 (2005) 6982 70
2001; Twitchett et al., 2001 among others). Many of
these events are still unresolved in most of the world
mainly due to an incomplete sedimentary record, the
particular lithological characteristic and the lack of
adequate fossils.
Over the last decade, several new techniques have
been developed to address this question. These meth-
ods show a large spectrum of possibilities for eluci-
dating the main factors responsible for the general
changes that occurred during this period of time,
including terrestrial and extraterrestrial causes.
Among the proposed terrestrial causes is a chain of
general changes occurring in response to severe mod-
ifications in the atmosphere as a result of the rapid
eruption represented by the Siberian traps basalt
floods (Renne et al., 1995; Kozur, 1998; Wignall,
2001). As a consequence, the collapse of terrestrial
ecosystems could have initiated a worldwide fungal
event at the top of the Permian, as suggested by Eshet
et al. (1995), Visscher et al. (1996) and Steiner et al.
(2003).
The present paper is based on a detailed study of a
spectacular specimen of Thuringian (Late Permian)
age consisting of a well-preserved fossil trunk show-
ing signs of interaction with fungi and found in a
middle zone of the Landete Formation, Iberian
Ranges, in the east of Spain (Fig. 1). In the Iberian
Ranges, the succession of Permian and Triassic sedi-
ments is represented by classic siliciclastic red-bed
deposits of continental origin (Sopena et al., 1988;
Lo pez-Go mez et al., 2002). The Late Permian sedi-
ments of the SE Iberian Ranges have been well-
known since the middle 20th century (Richter and
Teichmu ller, 1933; Boulouard and Viallard, 1971;
Viallard, 1973; Virgili et al., 1983, 1980; Lo pez-
Go mez and Arche, 1993; Arche and Lo pez-Go mez,
1999), however, until now, few well-preserved floral
elements have been found in these sediments.
Many publications have dealt with fossil woods of
different age, but few specimens have appeared so
close to the Permian-Triassic boundary (PTB) as the
trunk described here. There are also few descriptions
of fungalplant interactions in the Late Permian (Tay-
lor et al., 1994; Visscher et al., 1996; Taylor and
Taylor, 1997; Steiner et al., 2003, among others),
and only exceptional references to fossil woods bio-
degraded by saprophytic fungi (Stubblefield et al.,
1985; Taylor and Stubblefield, 1987; Stubblefield
and Taylor, 1988; Taylor, 1990; Taylor and Osborn,
1992). The specimen described here is the first Per-
mian-Triassic wood of its kind described in Europe.
We present an anatomical and taphonomical
description of the fossil trunk, the possible signifi-
cance of its interaction with a fungus and the general
characteristics of the sedimentary environment. The
main aim of this study is to improve our current
understanding of the palaeoenvironmental and
palaeogeographical characteristics of this enigmatic
time interval, in a continental area not far from the
westernmost coast of a prograding Tethys Sea that
reached the Iberian Basin during the early Anisian
(Middle Triassic).
2. Geological setting and stratigraphy
The present-day Iberian Ranges are part of an
intracontinental fold chain in the east Iberian micro-
plate, formed by Cenozoic inversion of the intracra-
tonic Iberian Basin. This basin represented part of a
complex of rift systems that developed during the
Permian in central and western Europe (Fig. 1) and
underwent several synrift phases until the beginning
of the Jurassic (Sopena et al., 1988; Lo pez-Go mez
and Arche, 1993; Salas and Casas, 1993; Arche and
Lo pez-Go mez, 1996; Van Wees et al., 1998; Lo pez-
Go mez et al., 2002).
The fossil trunk was found 4 km south of Landete
village, in the southern area of the Iberian Ranges
(Fig. 1), where the Late Permian sedimentary record
comprises two formations of alluvial origin (Arche
and Lo pez-Go mez, 1999): the Boniches Conglomer-
ates and the Alcotas Siltstones and Sandstones, from
base to top, respectively. The trunk was found close to
the top of a fine-medium grained subarkosic sand-
stone body, approximately in the centre of the Alcotas
Formation (Fig. 2). The Alcotas Formation crops out
in a large area of the Iberian Ranges, with generally
good outcrops normally related to anticline cores. This
unit lies conformably on the Boniches Formation or
unconformably on the Ordovician quartzites or slates
and is up to 125 m in thickness. In the upper contact,
the Canizar Formation overlies the Landete Formation
by means of a sharp contact that probably represents a
hiatus or mild unconformity (Lo pez-Go mez and
Arche, 1993).
C. Dieguez, J. Lopez-Go mez / Palaeogeography, Palaeoclimatology, Palaeoecology 229 (2005) 6982 71
2.1. The age of the fossil trunk and the Alcotas
Formation
Discerning the age of Permian sediments repre-
sented by siliciclastic red beds has always been pro-
blematic. For example, the Karoo Basin (McLeod et
al., 2000) is one of only a few examples in the world
that permit timing the evolution of fossil groups in red
beds by isotope contrasting. It is even more difficult to
find sections to compare the biotic evolution of marine
and terrestrial realms in the same area, as in the case of
some Jameson Land series, East Greenland (Twitchett
Fig. 2. Detailed stratigraphical section of the Landete Formation in the Landete area and location of the fossilised trunk.
C. Dieguez, J. Lo pez-Gomez / Palaeogeography, Palaeoclimatology, Palaeoecology 229 (2005) 6982 72
et al., 2001; Stemmerick et al., 2001). A further limita-
tion is that, in most of the world, the Permian-Triassic
transition shows substantial sediment gaps.
Only a few papers have described the Late Permian-
Triassic chronostratigraphy of the Iberian Ranges in
detail (Boulouard and Viallard, 1971; Sopen a et al.,
1988, 1995; Doubinger et al., 1990; Lo pez-Go mez et
al., 1998, 2002, among others), yet more remarkable is
that the Scythian (Lower Triassic) was never dated in
the Iberian Peninsula. Indeed, the incompleteness of
the Permian-Triassic fossil record is an interesting
matter of discussion (see Twitchett, 2001). The avail-
able chronostratigraphical data were mainly derived
from palynological assemblages, and many of these
assemblages appear in the Landete Formation (Bou-
louard and Viallard, 1971; Arche et al., 1983; Virgili et
al., 1983; Lo pez-Go mez and Arche, 1986; Doubinger
et al., 1990) or its lateral equivalent formations
(Ramos and Doubinger, 1979; Sopen a et al., 1995).
This would indicate a Thuringian (Late Permian) age
for this unit based on the presence of: Nuskoisporites
dulhuntyi, Lueckisporites virkkiae, Paravesicaespora
splendens and Klausipollenites schaubergeri.
The trunk was found 65 m from the base of the
section (Fig. 2), in which three palynological assem-
blages at 25 m, 59 m, and 62 m, from base of section,
have been described (Boulouard and Viallard, 1971;
Doubinger et al., 1990). The fossil trunk is therefore
considered Thuringian in age. Other trunk remains,
less than 80 cm long were found dispersed along the
base of the same level.
Palynological assemblages of both underlying
(Boniches Conglomerates) and overlying (Canizar
Sandstones) formations of the Landete Formation indi-
cate Thuringian (Late Permian) and Anisian (Middle
Triassic) ages, respectively (Doubinger et al., 1990).
However, we have to consider that the underlying
Boniches Formation association also shows Autunian
forms in its uppermost part, such as Vittatina and
Potoniesporites, indicating an early Thuringian age,
and that the assemblage of the overlying Canizar For-
mation was also obtained at the top of this formation.
2.2. Sedimentary characteristics and depositional
environment
The Alcotas Formation is made up of red siltstones
and clays, and associated lenticular sandstones,
including conglomerate bodies. The maximum thick-
ness is 170 m. Sandstones are arkoses, their clay
fraction consisting mainly of illite (Alonso-Azcarate
et al., 1997). Total mineralogy shows a substantial
punctual increase in hematite and dolomite in the
upper third of the formation (Fig. 2). Sandstone and
conglomerate bodies in the formation consist of
upward-thinning and -fining sequences, generally
less than 1 m thick. When well developed, as in the
section in which the fossil trunk was found, these
bodies show concave-upwards bases and do not gen-
erally exceed 550 m in lateral extension. Thinner
bodies have a somewhat flatter base and extend lat-
erally beyond several tens of metres. Palaeocurrents
towards the SE are persistently indicated.
The general sedimentary characteristics of the
bodies examined indicate that the Alcotas Formation
was deposited by sandy low-sinuosity braided rivers
isolated in floodplains that evolved to sandy mean-
dering and sheetflood distal braided deposits. The
sediments of the Landete Formation were deposited
in a basin consisting of a series of half-grabens with
the main sediment source located in the Iberian Massif
at the NW of the basin. These sediments were trans-
ported south eastwardly by axially orientated rivers
(Arche and Lo pez-Go mez, 1996).
Descriptions of both the sedimentological charac-
teristics of the Landete Formation and its sedimentary
environment are detailed in Lo pez-Go mez and Arche
(1993), Arche and Lo pez-Go mez (1996, 1999) and
Lo pez-Go mez et al. (2002).
During the deposition of most of the Alcotas For-
mation, the climate is believed to have been warm
and temperate, accompanied by a monsoon circula-
tion (Arche and Lo pez-Go mez, 1999). Although there
are obvious difficulties in comparing recent and
ancient climates, it should be noted that, according
to palaeomagnetic and palaeogeographic data, the
basin under study was located near the equator during
the Late Permian (Ziegler, 1988; Fluteau et al., 2001).
This hypothesis is also supported by the plant
remains found and the styles of fluvial sedimentation
of the Landete Formation indicating transport and
deposition by running water. On the other hand, an
increased hematite content in the middle part of the
studied section (Fig. 2), particularly abundant in red-
dish surface horizons in acidic sandy soils (Retallack,
2001), along with a decrease in the plant remnant
C. Dieguez, J. Lopez-Go mez / Palaeogeography, Palaeoclimatology, Palaeoecology 229 (2005) 6982 73
content of sediments, could tentatively indicate a
remarkable oxidation stage for this period of deposi-
tion, although other already existing factors could
also have been at work.
3. Anatomy and taxonomy of the fossil trunk
The fossil log is a very well-preserved silicified
specimen that was first described in Dieguez and
Lo pez-Go mez (1999). Its present size is 6.2 m long,
1.1 m in diameter (Fig. 3). At least, signs of one in situ
original branch clearly related to this trunk were found
in the substratum sandstone level analysed here. The
specimen is housed in Museo de las Ciencias de
Castilla-La Mancha (Cuenca, Spain).
The fossil specimen is comprised of a decorticated
secondary xylem, with uniform and homoxylic wood
and no growth rings, showing clear signs of silica
permineralisation processes and is dark-brown
coloured.
3.1. Anatomical description
The anatomy of the specimen was established by
examining fourteen thin transverse, radial and tangen-
tial sections of the secondary xylem. These thin sec-
tions were made using the conventional cutting and
polishing methods for optical microscopical study
being deposited in the Museo Nacional de Ciencias
Naturales-CSIC (Madrid, Spain).
Cross-section (Figs. 4a and 5ae). Growth rings
absent. Xylem rays uniseriate, about 25 Am width,
separated by a variable number of files of tracheids,
from 2 to 7 (average 3). Tracheids thin-walled, about
25 `m width, of variable shape (from isodiametric to
oval or polygonal in transverse section) depending on
compression and the proximity to other tracheids,
from 87.5 to 125 Am in size and arranged in regular
radial files. Neither the resin ducts nor the bordered
pits on the end wall of tracheids have been observed.
Nevertheless, there are pairs of bordered pits in the
wall of tracheids with torus located in the middle of
the membrane.
Tangential section (Figs. 4b, c, g and 5f). Rays
are mostly uniseriate ranging from 4 to 18 cells in
height. The upper and lower ends of the rays are
suddenly tapering which is a typical araucariaceae
structure. Ray cells thin-walled (about 12.5 Am)
varying in size in the same ray from 12.5 to 62.5
Am in diameter. Tangential bordered pits present at
the end of tracheids.
Radial section (Fig. 4df). Tracheids showed a
large, circular lumen with absence of spiral thickening
in their radial walls. Bordered pits in radial walls
(from 12 to 39) oval to rounded, about 40 Am in
diameter, arranged usually in one file (rarely two)
according to an araucaroid pattern. If uniseriate they
are arranged contiguously. When biseriate, they are
arranged alternately and become uniserate at the ends
of tracheids.
3.2. Taxonomy
Based on these general anatomical features, we
were able to classify the wood specimen as a Dadox-
ylon sp. of the Coniferophyta division comprised of a
Fig. 3. The studied fossilised trunk in the Landete section outcrop.
Hammer in the middle part of the trunk for scale. See also Fig. 2 for
the stratigraphical location of the trunk in the section.
C. Dieguez, J. Lo pez-Gomez / Palaeogeography, Palaeoclimatology, Palaeoecology 229 (2005) 6982 74
a b c
d e f g
100 m 100 m 100 m
100 m 100 m 100 m 100 m
Fig. 4. (a) Uniserate xylem rays and tracheids arranged in regular radial files, 31786, TS. (b) Short linear uniseriate rays cells, MNCN343,TLS.
(c) Long uniseriate rays and tangential bordered pits, MNCN342, TLS. (d) Uniseriate araucaroid bordered pits arranged contigously, 35078,
RLS. (e) Uniseriate bordered pits at the end of tracheids, 31874, RLS. (f) Biseriate bordered pits arranged alternately, 31874, RLS. (g) Ends of
rays tapering suddenly and tangential bordered pits (arrow), MNCN342, TLS.
Fig. 5. (a) Contiguous walls of two tracheids showing bordered pits with torus in central position (arrow) and diverse stages of delignification
31786, TS. (b) Lignin decay with separation of walls layers due to the fungus attack in an early stage of lignin removal (arrows), 31786, TS. (c)
Advanced stage of delignification. Only some zones of the tracheids walls and persistent wall corners can be observed. Tracheids are perceptible
as impressions except in the central area. (d) Early stage of decay. Note the different degree of decay in the different zones of wood, 31786, TS.
(e) Later stage of wood-decay. Persistent wall corners and middle lamellae of thacheids located close to a decay pocket, 31786, TS. (f) Early
stage of delignification of tracheids with zones without tangential walls (arrows) MNCN342, TLS.
C. Dieguez, J. Lopez-Go mez / Palaeogeography, Palaeoclimatology, Palaeoecology 229 (2005) 6982 75
single, homogeneous wood type (homoxylic). The
anatomy of the specimen closely resembles the sec-
ondary xylem of the extant Araucaria brasiliana Rich
(Araucariaceae).
The general anatomical characteristics of the trunk,
including homoxylic wood and bordered pits of the
araucaroid type, are consistent with those presented by
the genus Dadoxylon Endlicher and Araucarioxylon
Krauss. Literature on gymnosperm fossil wood
records a comprehensive discussion on nomenclature
of these two genera and diverse criteria for the taxo-
nomic attribution of specimens have been established.
These criteria were based on different aspects viz.: age
of the specimens (Seward, 1917; Stewart, 1983), pre-
sence of uniseriate or multiseriate rays (Maheshwari,
1972), presence of pith and primary xylem (Lepe-
khina, 1972), and presence/absence of araucaroids
remains in the same horizon from which the specimen
of wood comes (Seward, 1917). More recently, new
nomenclatural revisions of the homoxylous woods
have been made (Philippe, 1993; Philippe et al.,
1999; Bamford and Philippe, 2001) mostly based on
Mesozoic material.
Following the criteria established by Seward
(1917, p. 249) and Stewart (1983, p. 344), the speci-
men has to be ascribed to the genus Dadoxylon End-
licher. According to these criteria, wood specimens
showing araucaroid bordered pits collected from
Palaeozoic sediments should be denoted as such. In
addition, Seward (1917, p. 250) advocated the assign-
ment to Dadoxylon for those fossil wood specimens
collected in a horizon in which unmistakable repre-
sentatives of the Araucariaceae are lacking. The fossil
record of Dadoxylon is difficult to establish since
different systematic fossil classification systems have
been applied such that some Dadoxylon trunks have
been attributed to Araucaroxylon, or vice versa. The
Dadoxylon record and related palaeoxylologic studies
for the Iberian Peninsula are restricted to citations by
Broutin (1978) for the Lower Permian of R o Viar
(Seville, Spain), and by Garc a Gimenez et al. (1983)
for the Mesozoic of Palmaces de Jadraque (Guadala-
jara, Spain).
3.3. Remarks
The specimens secondary xylem shows a replace-
ment by silica in fossilisation processes. Although
preservation is exceptional, in some areas the com-
plete destruction of the cell walls of tracheids can be
observed (Fig. 5bf). These areas are associated with
other zones of the wood in which cell dissociation in
the tracheids is patent. This type of wood decay (Fig.
5b) is extraordinarily similar to that shown by speci-
mens of Araucaroxylon from the Permian and Triassic
of Antarctica (Taylor and Stubblefield, 1987, Fig. 2;
Stubblefield and Taylor, 1988, Fig. 2f).
In addition to this decay, the wood also contains
spaces (pockets) lacking conductive elements sur-
rounded by cells with secondary walls clearly dis-
torted by lignin decay, also comparable to those
described by Stubblefield and Taylor (1988, Fig. 2f),
and pockets surrounded by tracheids, modified in
shape, at different stages of lignin decay (Fig. 5b,
c). These patterns of decay in the secondary wood
can be attributed to the actions of saprophytic fungi as
described by Taylor and Osborn (1992). This latter
work (Fig. 7 thereof) also describes a Permian speci-
men of Araucaroxylon with cell corners persisting in
tracheids after delignification by fungi. Similar results
have been observed in our specimen (Fig. 5c, e).
Saprophytism of wood is very common today, but
the fossil record of this interaction between plants and
fungi, known as wood rot, is extraordinarily scarce,
and the few existent references almost exclusively
relate to trunks of similar age to the specimen
described here (Taylor and Stubblefield, 1987; Stub-
blefield and Taylor, 1988; Taylor, 1990; Taylor and
Osborn, 1992).
Recent analysis of the effects of fungal infection of
plant remains (conducted on a Cenomanian conifer)
indicate that distorted cell walls show substantial
amounts of hydroxysuccinic acid and functionalised
benzoic compounds that were interpreted as degra-
dation products of lignin by fungi (Nguyen Tu et al.,
2000). The decomposition of organic matter by micro-
bial or fungal action may have also contributed to
significant amounts of polysaccharides found in Late
Permian sediments in the Dolomites in northern Italy
(Sephton et al., 1999). The findings of both these
studies indicate that this fungal activity could be
more intense than observed in more ancient sediments.
The activity of saprophytic fungi, particularly that
of basidiomycetes, leads to lignin degradation, trans-
forming organic C to inorganic C by an enzyme action
known as wood rot (white rot and white pocket rot).
C. Dieguez, J. Lo pez-Gomez / Palaeogeography, Palaeoclimatology, Palaeoecology 229 (2005) 6982 76
Ligninolytic enzymes have a broad specificity and
decay processes occur in an obligate aerobic environ-
ment (Pointing, 2001). Initially the fungus focuses its
enzyme activities on the middle lamella, which sepa-
rates the woods cells leading to their final destruction
(Fig. 5af). This action affects cellulose and the lignin
which makes up the tracheids. Lignin biodegradation
mechanisms in living material have been mainly
explored by evaluating the activity of basidiomycete
fungi (Leonowicz et al., 1999; Steffen et al., 2000;
Papinutti and Forchiassin, 2000). There are, however,
some recent studies on fossil material (Nguyen Tu et
al., 2000) from Cretaceous conifers that describe the
action of ascomycete fungi that act on the lipid frac-
tion. Further research efforts have explored potential
factors affecting the ligninolytic activity of saprophy-
tic fungi (Godio et al., 2000) and their action on
organopollutants of similar chemical structure to lig-
nin (Pointing, 2001).
Analogous decay patterns have been observed in
fossil and living wood, suggesting a similar decay
mechanism at the biochemical level (Stubblefield
and Taylor, 1986). Thus, correlations may be made
among the mechanisms and factors affecting the
extant wood and those that affected the fossils. Lignin
degradation is mainly attributable to basidiomycetes
but there are also a few ascomycete and deuteromy-
cete fungi capable of wood rot decay. Based on the
anatomical alterations and type of the key pattern
observed, we attribute this decay to the action of
saprophytic fungi. However, given this indirect evi-
dence and the lack of fossilised fungal remains such as
mycelia, hyphae or spores, it is not possible to identify
the agent producing the decay.
4. Taphonomy of the fossil trunk
4.1. Biostratinomic processes (transport and abrasion)
After the death of the tree and fall of the trunk,
necrolytic processes may have been induced by
fungi, given the appropriate humid and temperate con-
ditions of the area for their proliferation, probably
along with high CO
2
concentrations (Fluteau et al.,
2001; Retallack, 2001; Ku rschner, 2001). High con-
centrations of atmospheric CO
2
would stimulate the
trees physiology, development and growth, by enhan-
cing leaf photosynthesis, suppressing plant respiration
and reducing transpiration. Biomass production, litter-
fall would also be enhanced, as microbial biomass
proliferation (especially fungi) and the activity of
decomposers such as fungi increased (Ceulemans et
al., 1999; Osborne and Beerling, 2002). The quality of
preservation of areas unaffected by saprophytic fungi,
indicate the trunk was not substantially affected by
further biotic activity (of bacteria and/or fungi). Bear-
ing this in mind along with the decomposition time of a
trunk (Martin, 1999, Table 5.3), we could argue that
fungal infection began very soon after the fall of the
tree.
Although the extent of fungal infection cannot be
determined in the specimen studied and consequently
the decay quantified, it is evident that, in larger or
smaller measure, the physical characteristics of the
wood would be affected by the disappearance of lignin.
In addition, altering the anatomy of the wood, the
action of wood rotting fungi also modifies the physical
properties of the wood, e.g. their water-holding capa-
city (Benito, 1943; Boddy, 1986; Norden and Paltto,
2001).
Sedimentological data indicate general low energy
shallow water fluvial systems during the deposition of
the trunk (Arche and Lo pez-Go mez, 1999). It is very
possible that the channel capacity of these fluvial
systems was insufficient to transport the trunk over
a long distance and it is likely that these channels were
abandoned very soon.
There are no indications of further modification to
the physical characteristics of the trunk such as resis-
tance to compression, flexibility and hardness and
stiffness, with the exception of the absence of defor-
mation or breakage, from which we could infer that
the trunk was not severely altered.
4.2. Fossil diagenetic processes
Diagenetic analysis of the fossil trunk indicates
several linked stages that clearly started as soon as
the trunk became buried. Its relatively well-preserved
internal details could indicate that decay was early
inhibited and that mineralization progressed rapidly
due to fluvial dynamics inducing the quick burial of
the trunk. Similar cases were described by Fielding
and Alexander (2001) in Australia for recent and
Permian fossil trees. The rapid subsidence of the
C. Dieguez, J. Lopez-Go mez / Palaeogeography, Palaeoclimatology, Palaeoecology 229 (2005) 6982 77
Iberian Basin during Thuringian times has also been
reported (Salas and Casas, 1993; Arche and Lo pez-
Go mez, 1996; Van Wees et al., 1998). Mineralization
is largely dependent on the depositional environ-
ment, although the precise chemistry of this process
is not well understood. It seems that partial decay is
needed for silica preservation, and early diagenetic
silicification is also a prerequisite for exceptional cases
of microorganism preservation. However, further ad-
vanced decay would have caused the complete or
nearly complete destruction of internal details except
for those comprised of more complex macromolecules
(Allison, 1988; Allison and Briggs, 1991).
Two quartz forms were differentiated by petrogra-
phical analysis of thin sections: megaquartz and
microquartz (Fig. 6a,b). The microcrystalline mor-
phology of the latter indicates the replacement of
original chalcedonic quartz. Both megaquartz and
microcrystalline quartz infill pore spaces, while gen-
eral diagenetic modifications give rise to megaquartz
through dissolutionreprecipitation reactions.
The outer area of the secondary xylem of the
trunks internal structures were the best preserved,
since penetration of liquid silica probably com-
menced in this region. The logs inner areas were
therefore progressively partly sealed from further
penetration, and decay probably progressed leading
to destruction and to the formation of cavities filled
with crystalline quartz.
Fluid migration in a generally acid medium prob-
ably conditioned the replacement of the original ele-
ments of the plant cell structure by microquartz in an
initial stage of diagenesis. This first fluid migration
stage could have formed the red-brown iron oxide
coating in the spaces between the walls of the trac-
heids, wherever iron was readily available in stream
water. A later stage is indicated by the appearance of
microquartz, probably both by replacement and pore-
infill processes, as suggested by the straight bound-
aries of the crystals or triple points where growth
meets, which are highly characteristic of pore-filling
cements (Adams et al., 1984). Macroquartz crystals
could indicate a subsequent diagenetic stage. This
form of quartz newly occurred in the xylem rays,
where recrystallisation was still possible.
Crystal growth in microfractures of the specimen is
also suggestive of several stages. An early stage prob-
ably involved refill of microquartz in synsedimentary
microfractures that were probably formed during the
readjustment of the trunk during the early stages of
burial. Iron oxide was also probably deposited on the
microfracture walls during these processes, while
silica would have migrated along these microfrac-
tures. Microfractures that cross-organic structures
show a clear later stage of recrystallization. These
newly generated microfractures would be refilled
after the migration of silica-saturated fluids once
again via the newly created free spaces.
Polycrystalline quartz with crystals elongated in
preferred orientations and sutured boundaries between
crystals were relatively common in the fossilised
trunk. These probably appeared as a result of stress
Fig. 6. (a) Micro- and macroquartz crystals showing preferential
fluid migration routes controlled by the cell organisation of the
fossil trunk. (b) Secondary macroquartz crystals of triangular
shape (top left) and perpendicular veins reflecting a later stage of
crystal growth (lower right).
C. Dieguez, J. Lo pez-Gomez / Palaeogeography, Palaeoclimatology, Palaeoecology 229 (2005) 6982 78
conditions due to changes in pressure and temperature
during several later phases of the Alpine Orogeny
(Salas and Casas, 1993) and the appearance of micro-
folds and microfractures could have promoted fluid
circulation. In contrast, the frequent finding of a well-
preserved cell structure suggests a limited degree of
compaction or its prevention in part by early miner-
alisation, such that tissue permeation could take place
(Carson, 1991).
Although there are several possible sources of
silica through geologic time (see Carson, 1991), in
the case of the particular lithology of the Landete
Formation, some of the silica may have been derived
from detrital quartz, as well as from the transforma-
tion of the clay minerals themselves, such as the
alteration of montmorillonite to kaolinite during sub-
aerial weathering (Alonso-Azcarate et al., 1997).
5. Discussion
The Permo-Triassic boundary marks a period in
which known species numbers showed a fall of
around 20% and in which the sedimentation of
plant-bearing rocks was low, at least in Europe
and North America (Niklas et al., 1983). The Late
Permian saw the adaptive radiation of the seed
plants that culminated in the gymnosperm-domi-
nated Mesozoic.
Known stratigraphic ranges of natural conifer taxa
recognised in Late Permian flora assemblages from
western, central and southern Europe, such as Wal-
chiaceae or Ullmanniaceae, do not cross the PTB.
Species of these Late Permian families produced
pollen grains corresponding to several palynological
species such as Jugasporites, Lueckisporites, or Nus-
coisporites dulhuntyi in the case of species of Orti-
seia. These were a prominent component of the
xerophilous Late Permian flora of the southern Alps
(Poort et al., 1997). In a vertical interval of less than
5 m, the Landete Formation shows some of the last
Permian levels known to date in Iberia. These levels
also contain the conifers Dadoxylon sp. and Ullman-
nia sp. and the palynological species L. virkkiae and
Nuscoisporites sp. (Doubinger et al., 1990).
Conifer dieback may have provided the organic
resource supply for fungal proliferation and has been
well described in Europe, North America, Asia,
Africa and Australia for the Late Permian (Eshet et
al., 1995; Visscher et al., 1996; Looy et al., 2001;
Steiner et al., 2003), where fungi probably adapted
and responded quickly to the environmental distur-
bance. Thus, the dieback of arboreal vegetation may
have been a worldwide event that affected the terres-
trial biosphere, irrespective of the local nature of
climatic and flora changes (Poort et al., 1997; Looy
et al., 1999). In several sections of the Southern Alps
(Visscher and Brugman, 1986; Cirilli et al., 1998;
Visscher et al., 2001), Israel (Eshet et al., 1995)
and South Africa (Steiner et al., 2003), there are
reports of the disappearance of the gymnosperm-
dominated palynoflora of the Late Permian L. virk-
kiae from a claystone horizon almost exclusively
containing abundant fungal remains and carbonised
terrestrial plant debris. We do not exclude the possi-
bility that the above-mentioned 5 m interval of the
upper Landete Formation corresponds to part of a
large-scale loss of standing biomass in the western-
most Tethys as one of the stages in the worldwide
biotic crisis of the Late Permian.
If adverse conditions at the end of the Permian,
such as acidification, anoxia or any other cause or
causes (see Kozur, 1998; Wignall and Hallam, 1992;
Erwin, 1993; Hallam, 1994) that altered the develop-
ment of plants, allowing the opportunistic prolifera-
tion of fungi, this competition must have also
conditioned the different plant groups. If the cause
or causes suggested above led to a general rise in
atmospheric CO
2
levels (Yin and Tong, 1998;
Kozur, 1998), the global vegetation mass would initi-
ally increase leading to reduced soil nutrients.
6. Conclusions
We report the discovery of an extremely well-pre-
served fossilised trunk (6.2 m long, 1.1 m wide) of
Thuringian (Late Permian) age in the Landete Forma-
tion, SE Iberian Ranges, Spain. The general anatomi-
cal features of the fossil correspond to those shown by
the genus Dadoxylon Endlicher of the Coniferophyta
Division and are similar to those of the present-day A.
brasiliana Rich (Araucariaceae).
Detailed petrographical study permits to observe
different processes such as decay processes due to
fungal activity. These processes include the destruc-
C. Dieguez, J. Lopez-Go mez / Palaeogeography, Palaeoclimatology, Palaeoecology 229 (2005) 6982 79
tion of lignin, indicating the activity of saprophytic
fungi, an interaction previously described for the Per-
mian of Antarctica. These processes could also be
related to those worldwide fungal events at the top
of the Permian as a consequence of the collapse of
terrestrial ecosystems and the increase of organic
material proceeding of the dieback of forests.
Sedimentological approaches indicate that fluvial
system was insufficient to transport the trunk over a
long distance. Fossil diagenesis processes indicate
early dissolutionreprecipitation reactions and differ-
ent stages of recrystallisation.
Acknowledgements
This paper is a contribution to PICG Projects 458,
465 and PB98-0488 and BTE2002-00775 of the
Spanish Ministry of Science and Technology. We
thank Gilberto Herrero for preparing the thin sections.
Helpful reviews by Sharon Klavins and an anon-
ymous referee improved the content of the article.
Special thanks are extended to B.K. Ferguson, A.
Arche and F. Surlyk for constructive comments. We
also thank Ana Burton for the revision of the first
English version.
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