Cour. Forsch.-Inst. Senckenberg
256
291–304
4 Figs, 1 Tab.
Frankfurt a. M., 15. 11. 2006
A century of research on the classical locality of Tegelen
(province of Limburg, The Netherlands)
With 4 figs, 1 tab.
Lars W. VAN DEN HOEK OSTENDE & John DE VOS
Abstract
In 1904 EUGÈNE DUBOIS published the first report on fossil mammals from the vicinity of Tegelen. After a
century of research the clay-pits from the area are known as a classical locality for Villafranchian mammals. ANTJE SCHREUDER’s work was particularly important for making the fossils from Tegelen known. Her
papers, however, also indicate some problems. The large mammal fossils were gathered by clay-workers
in the local ceramic industry, and the fragmentary nature of the fossils is partly caused by injudicious handling while extracting the bones from the clay. Because the Tegelen fossils are chance finds rather than the
results of careful excavations, their exact provenance is not established. The only exception is the collection
of micromammals, which was gathered by THIJS FREUDENTHAL during expeditions in the 1970s. Large size
variation in the cats and hyenas, and the presumed presence of Stephanorhinus kirchbergensis cast doubt
on to whether the fossils from Tegelen represent a single time frame.
Key words:
Tegelen, Villafranchian, fossil mammals, faunal list
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Introduction
The locality is the most important entity in mammal palaeontology. A species is a concept, subject to the interpretation of scholars, and can often change as the available
material is revised. A locality, in contrast, is real. It is a
place where you can get your hands dirty while unearthing fossil bones. To the palaeontologist it also represents
a window to the past, a stage in evolution, allowing a
glimpse to a particular time frame.
By placing the localities of a particular area in
stratigraphical order, we obtain a picture of the regional
development. Comparing localities from different areas,
particularly if supplemented with radiometric dates and/or
palaeomagnetic sequences, we can come to interregional
correlations. Thousands of localities of different time
frames are known all over the world by now. Ideally, a
palaeontologist should take as many localities as possible
into account when reconstructing the history of life. This
raises, however, many practical problems, particularly
when dealing with localities outside ones area of study.
Therefore, interregional correlations are often based on the
comparison with classical localities. This is, for instance,
the basis for the subdivision of the Neogene and Paleogene into MN-zones (MEIN 1975, DE BRUIJN et al. 1992)
and MP-zones (SCHMIDT-KITTLER 1987), respectively.
The reference localities for these zones are the ones best
studied and most cited in the literature, and which are
believed to be best known to all the scholars interested in
that particular time frame.
There are clear advantages in choosing well-known localities in schemes for interregional correlations. However,
there are drawbacks as well. Many of these classical localities were excavated in earlier times without the scrutiny
and sophistication used in modern excavations. Material
from neighbouring sites, but from different levels, may
have been entered in museum collections under the same
locality name. Herein lies the importance of re-excavating
Authors’ address: Dr. Lars VAN DEN HOEK OSTENDE & Dr. John DE VOS, Nationaal Natuurhistorisch Museum Naturalis,
P.O. Box 9517, 2300 RA Leiden, The Netherlands, <Hoek@Naturalis.nl>, <Vos@Naturalis.nl>
© E. Schweizerbart’sche Verlagsbuchhandlung (Nägele u. Obermiller), 2006, ISSN 0341-4116
VAN DEN
HOEK OSTENDE & DE VOS: A century of research on the classical locality of Tegelen
classical sites such as Senèze (DELSON et al. 2004) and St.
Vallier (DEBARD et al. 1994, VALLI 2001). However, many
classical localities are no longer accessible or have been
exhausted. They can still be studied from the material
saved in (museum) collections. In these cases knowing
the history of the collection is essential. The method of
excavation, the material’s original state of preservation,
the way of processing the finds, and even what has happened with the material after it has been stored, they can
all influence our interpretation of a fossil collection.
In this article we discuss such a classical locality that
is no longer available for further excavations. The clay
pits near the Dutch village of Tegelen (province of Limburg) are known for over a century to yield fossil remains
of mammals (DUBOIS 1904). When REID & REID (1915)
established the Tiglian on the basis of the seed flora from
the locality, the pits became a type locality for a chronostratigraphic unit that is still widely used. Its reputation as a
classical locality was strengthened when, next to the large
mammals and seeds, also the pollen (ZAGWIJN 1963) and
micromammals (FREUDENTHAL et al. 1976) were studied.
Although the Tegelen fauna has been the subject of numerous studies over the years (e.g. REUMER 1984, SPAAN 1992,
TESAKOV 1998, REUMER & VAN DEN HOEK OSTENDE 2003,
VAN DEN HOEK OSTENDE 2003, O’REGAN & TURNER 2004),
no recent overview of the entire fauna has been given,
with the exception of an exhibition guide (VAN DEN HOEK
OSTENDE 1990) and a popular article (VERVOORT-KERKHOFF
& VAN KOLFSCHOTEN 1987), both in Dutch and therefore
not widely accessible to the scientific community.
In this paper we give a review of the results of a hundred years of palaeontological research in the Tegelen
clay pits, to show how we reached our present state of
knowledge. The history of the research has been extensively dealt with elsewhere (ZAGWIJN 1998, VAN DEN HOEK
OSTENDE 2004), but since knowledge of the history of the
collections is vital for understanding the fauna, it is shortly
reiterated here.
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The Tegelen Clay
The clay-beds near Tegelen are known since Roman times.
The village even owns its name to the Latin word for tile,
tegula. Tegelen had a flourishing ceramic industry. Till the
end of the 19th century its main products were household
pottery, but as the pottery shops were out-competed by
cheaper alternatives, the industry shifted to the production
of tiles and (chimney) bricks. Because the clay from the
Tegelen region proved very suitable for these high-quality products, it was extensively exploited. During these
excavations, which in the first part of the 20th century were
done by manual labour, bones were regularly encountered
by the clay-workers. Hence, Tegelen owns its reputation
1
as a fossil mammal locality to the exploitation for the
ceramic industry.
Not all of the clay-beds in the region are of the same
age. A bluish clay with intercalated lignite beds outcrops
near Reuver, a few kilometres to the south of Tegelen.
Based on the seed flora, REID & REID (1915) showed that
this clay is clearly older than that near Tegelen. The Reuver
Clay lies in the upper part of the Kieseloolite Formation,
which is constituted by Upper Pliocene fluvial sediments.
The Kieseloolite Formation is overlain by the recently
introduced Waalre Formation, which includes the former
Tegelen Formation together with the ‘Rhine’part of the
former Kedichem Formation (WEERTS et al. 2003). The
Waalre Formation, which in the vicinity of Tegelen consists
of fluvial sediments, was deposited in the Late Pliocene1
and Early Pleistocene by the Rhine-Meuse system. The
basis of the formation in the region consists of gravels and
sands, while it is capped by clay layers that seem to be a
mixture of channel fills and laterally extending floodplain
clays (WESTERHOFF et al. 1998). These clays are certainly
not all of the same age. The exposures in the Maalbeek
pit were shown to be older than those in the former pits
in the direct vicinity of Tegelen (WESTERHOFF et al. 1998),
an age difference that had already been assumed from the
presence of tapir and mastodont remains in the pit. An
array of northwest to southeast-ranging major and minor
faults complicates the geology of the region (fig. 1). One
of these structures, called the Tegelen fault, runs between
the Maalbeek pit and the other Tegelen pits. Also, the
clay beds in the pits near Tegelen itself belong to different sedimentary cycles. This is most apparent in the pit
Russel-Tiglia-Egypte, where a stream gully cuts through
the underlying clay beds. Although KORTENBOUT VAN DER
SLUIJS & ZAGWIJN (1962: 32) admitted that “a certain stratigraphic gap between the two cycles may be present”, it is
clear that they did not consider this gap to be very large.
The basis of the stream gully infilling consists of sandy
clay with a high amount of plant remains. According to
KORTENBOUT VAN DER SLUIJS & ZAGWIJN (1962), similar
sandy beds were found at the base of the clay in the
Canoy-Herfskens pits, formerly called the Russel-Tigliapit, and the Russell-Tiglia-Wambach-pit, indicating that
the clay in these pits would be of the same age as the gully
infilling in the Russel-Tiglia-Egypte pit. This, of course,
is a very rough correlation of the clay beds and should be
regarded as a preliminary assessment. KORTENBOUT VAN
DER SLUIJS intended to study the various clay-beds for his
PhD-thesis, which, however, never appeared. That the
method is prone to error is shown by the correlation of
the Maalbeek pit, the sediments of which KORTENBOUT
VAN DER SLUIJS & ZAGWIJN (1962) believed to be younger
than those in the Russel-Tiglia-Egypte pit, whereas they
are in fact older (WESTERHOFF et al. 1998). Nevertheless,
KORTENBOUT VAN DER SLUIJS & ZAGWIJN’s (1962) descrip-
The Plio-Pleistocene boundary used in this article follows the GTS. This may account for some differences with particularly Dutch literature, in
which this boundary is placed earlier.
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Cour. Forsch.-Inst. Senckenberg, 256, 2006
Fig. 1: The location of the various clay pits near Tegelen (reproduced from KORTENBOUT VAN DER SLUIJS & ZAGWIJN 1962).
tions clearly show that the beds near Tegelen belong to at
least two sedimentary cycles.
Fig. 2: Much of our knowledge on the Tegelen fauna and the
way it was excavated we owe to the work of ANTJE SCHREUDER
(1887–1952).
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Taphonomy
The great paradox of Tegelen is that although it is known as
a classic fossil mammal locality, no excavations were ever
organised (VAN DEN HOEK OSTENDE 2004). Instead, chance
finds of clay workers were gathered and palaeontologists
periodically collected these up during visits to the various
clay pits. Hence, all the Tegelen fossils are of uncertain
stratigraphic provenance. Most of our knowledge on how
the bones were gathered we owe to ANTJE SCHREUDER
(fig. 2). In her articles, particularly those written in Dutch,
she gave little anecdotes on how the fossils had been recovered. Thus, she related about the important role of the clayworkers, who even retrieved fossils of small mammals.
“Notwithstanding the fact that the clay-layers worked
35 year ago, when Dubois, Krause and Weingärtner
brought together the rich collections now preserved in
Haarlem (Teylers Museum), Berlin (Preuss. Geol. Anst.)
and in Amsterdam (Zoolog. Museum of the Univ.), we owe
it to the spade-digging by the firm of ‘Canoy-Herfkens’
that in the last decennium new mammals, such as Desmana
tegelensis, Hypolagus brachygnathus and Pannonictis pliocaenica have been recorded” (SCHREUDER 1945: 158).
The fossil material from Tegelen is very fragmented,
with only a few favourable exceptions. This seems mostly
due to preservation of the material and the fact that it was
pulled out of the clay by laymen.
“Quite undamaged specimens are rare in the collections, as in the clay they are wet and often so soft, that they
become injured by the spade of the workmen; they break
still more when pulled out of the clay. Moreover, when
dried, they are often so burst, that they must be glued immediately on the spot, or the loss of some fragments would
make a restoration impossible (SCHREUDER 1945: 158).”
“The bones of the elephant, when wet, are so little
resistant that the spade cleaves them without the workman
observing this” (SCHREUDER 1945: 158).
The point is well illustrated in SCHREUDER (1949) by
fossils of the hyena that were obtained by the Maastricht
museum in 1943, and by bear fossils which were collected
for the Leiden museum in 1949. In the case of the hyena,
a skull fragment had been found with part of the P4 associated with some isolated lower molars. The bear material consisted of some isolated upper and lower molars.
SCHREUDER (1949) assumed in both cases that the skull
and mandible must have been present, but were destroyed
during the extraction from the clay. She wrote:
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VAN DEN
HOEK OSTENDE & DE VOS: A century of research on the classical locality of Tegelen
“Collecting without expertise ever so often lead to
only the very dark and conspicuous teeth and molars being
noticed by the clay workers and them being – as it were
– plucked from the jawbone, and even then just some, and
certainly not the entire dentition. The fossil bone cannot be
distinguished that easily, and is furthermore soft, as long as
it is still in the clay. If one were to bring the clay workers
to collect such a group of molars and teeth together with
the embedding sediment, so bone and clay together, and
store it, one would certainly obtain considerable pieces of
the skull and mandible, provided the preparation would be
done in a laboratory. Preferably, of course, osteologists
should be found willing to come to Tegelen and to collect
there, because they know what can be expected in the direct
vicinity once a molar or bone has been found” (SCHREUDER
1949: 115, translated from Dutch).
And even if fossils were found, it was sometimes a
matter of easy gain, easy loss, as is clear from the first
finds of Leptobos in the clay.
“How little notion there is about the value of the fossils may be shown by the fact that we owe the first fossils
of the ox of Tegelen, viz. three upper molars, only to the
circumstance that the hole in the trousers of the finder, a
clay-worker, wasn’t larger still; all the smaller elements
of the dentition had fallen through on the way from the
pit to his home” (SCHREUDER 1936a: 207, translated from
Dutch).
Thus, it is clear that the conditions in which the fossils
were found were far from perfect. Another aspect that from
SCHREUDER’s articles is that the fossiliferous beds varied in
richness. It is ironical that during the period she did most
of her work, finds were relatively few.
“That the clay-layers now in working are much poorer
in fossils than those of former years is due to the fact that
the fossils are heaped up in a certain region of the deposit,
from which the working has shifted more and more. The
proprietors expect that within ten years the rich spots will
come into working again, and it is to be hoped that there
will be some scientific authority to prevent the destruction
of the majority of these highly valuable fossils, which most
of them have undergone during the clay digging of the past
years” (SCHREUDER 1945: 158–159).
These quotations show that later in life SCHREUDER
became more cynical about the way the collections were
formed, emphasising as she did the unprofessional way
in which the material was retrieved from the clay. This
certainly explains why such a large number of Tegelen in
the various collections is so fragmentary. After her death,
collecting continued in exactly the same fashion as before.
Probably, there were no alternatives. Although much material has been collected, the various Tegelen collections are
not very large, if one considers that they represent seventy
years of collecting, and that during that period tremendous
amounts of clay were excavated for the ceramic industry.
This probably reflects the fact that the clays were not so
very rich in fossils. From SCHREUDER’s papers we know that
concentrations did occur, but whether these would have
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294
been rich enough to organise a proper excavation is very
much open to doubt. Nevertheless, sizable collections are
preserved in Teylers Museum in Haarlem, in The Zoological Museum Amsterdam, in the Natuurhistorisch Museum
Maastricht, and in the Rijksmuseum voor Geologie en
Mineralogie (now Naturalis) in Leiden.
The collections
The Teylers collection, which was formed in the years
1904–1933, contains the first fossil mammalian finds from
the Tegelen clay-pits. These were collected by a young
medical student called LAURENS STIJNS. This amateur palaeontologist contacted EUGÈNE DUBOIS, whose fame was
rapidly rising after the find of Pithecanthropus in Java.
Among his other positions, DUBOIS was the curator of the
Paleontological-Mineralogical Cabinet of Teylers Museum
in Haarlem. DUBOIS was slow in responding to STIJNS’ first
request for assistance, but his interest grew after he visited
the clay-pits in 1902. He obtained STIJNS’ collection ‘for
study’ and set up a deal with one of the pit-owners, AUGUST
CANOY. Fossils found by the clay-workers were regularly
sent to Haarlem. The Teylers Museum provided handsome
compensation for the finds. Thus, DUBOIS, and later workers as well, never excavated bones himself, but simply
gathered the specimens incidentally found by others.
DUBOIS published very little on the Tegelen palaeontology. He introduced the locality, publishing the first faunal
list (DUBOIS 1904). A year later, DUBOIS (1905) described
the deer fossils. At the same time he enticed others to
work on Tegelen. The bear fossils (Ursus etruscus) were
described by NEWTON (1913). Several years before NEWTON
(1907) had also described the first micromammal from
Tegelen, Mimomys pliocaenicus, which had been found
while sieving the clay for fossil seeds. This study of the
seed flora was undertaken by CLEMENT REID, a palaeobotanist who had worked on the seed flora from the Cromer
Forest Beds. As DUBOIS (1904) claimed that the Tegelen
Clay was an equivalent of the Cromer Forest Beds, the
interest of REID in the flora was obvious. REID did not
concur with DUBOIS that Tegelen was coeval to the Cromer
Forest Beds. He published his finding together with his
wife ELEANOR (REID & REID 1915). They showed, on the
basis of the percentage of Tertiary floral elements, that
Tegelen is older than the British localities. Furthermore,
they also demonstrated that the clay just south of the village
of Tegelen contains a higher number of exotic elements,
indicating an even older age. DUBOIS (1904) had assumed
a similar age for all the clay beds in the region. REID &
REID (1915) created the Teglian (= Tiglian) for the clay
beds near Tegelen, and the Reuverian for the beds that
outcrop south of the village.
By the time REID & REID (1915) had created the Tiglian,
DUBOIS had lost interest in the Tegelen Clay beds. When
VAN REGTEREN ALTENA (1951) catalogued the Tegelen collection kept in Teylers, he found a box sent in 1913 that
Cour. Forsch.-Inst. Senckenberg, 256, 2006
had never been opened. Others continued collecting in the
clay pits. Some collections were sent to Germany (ZAGWIJN
1998), others to the Amsterdam museum and still others
to the mission museum in Steyl. The latter collection is
now part of the Tegelen collection in Naturalis. DUBOIS
did supervise two PhD theses on the Tegelen fauna. The
first was by a Jesuit priest called father BERNSEN, who
was DUBOIS’ assistant in the collection of ‘Indian fossils’
(from Java and Sumatra) in the Leiden museum. His
thesis dealt with the rhinoceros fossils from the Tegelen
clay (BERNSEN 1927). DUBOIS also held a position at the
University of Amsterdam and his assistant there, ANTJE
SCHREUDER, wrote her thesis on the beavers from Tegelen
(SCHREUDER 1928). After his thesis, BERNSEN set out to describe the entire Tegelen fauna in a series of articles in the
‘Natuurhistorisch Maandblad’ (BERNSEN 1930b, 1931a–d,
1932a, b, 1933/34). His untimely death prevented him
from finishing this task, and ANTJE SCHREUDER completed
the last paper in the series.
The Amsterdam and Maastricht collections were
formed in the years 1933–1950. After DUBOIS retired,
ANTJE SCHREUDER found herself without a formal position.
She was allowed to continue her studies at the Zoological
Museum in Amsterdam. Because of her fragile health, she
focused on the study of small mammals (VAN DET & VAN
DEN HOEK OSTENDE 2003). Her second interest was Tegelen, on which she published a series of papers (SCHREUDER
1928, 1929, 1935, 1936a, b, 1945, 1946, 1949). The Amsterdam museum already held a small collection of fossils
from the clay pits, which consisted of material collected
by father WEINGÄRTER between 1909 and 1912, and the
BÖHMERS and VAN BEMMEL collection, gathered between
1928 and 1934. SCHREUDER was recognised as a specialist
both in The Netherlands and abroad. New fossils were
therefore sent to the Amsterdam museum and formed a
small but important collection.
The collection in the Natuurhistorisch Museum
Maastricht was likely gathered in the same period. In the
catalogue entries date (but the years of acquisition are
registered only for a relatively small part of the collection) between 1930 and 1946. Many of the fossils were
determined by ANTJE SCHREUDER, which places them in
the same time frame.
Macro- and micromammals of the Leiden collection
in the years 1948–1977: In one of her last articles on the
Tegelen mammals, SCHREUDER (1949) ended with the
sentence “May ‘Leiden’ as well as ‘Groningen’ succeed
in adding valuable fossils to the Tegelen collections, now
that the clay layers in exploitation are again richer in fossils than they have been for years.” It was Leiden that took
up the challenge. However, as before no true excavations
were organised, but material was still collected from the
clay-workers during visits to the various pits. The trips
were initially organized by VAN DER VLERK and BROUWER.
VAN DER VLERK was building a stratigraphical framework
for The Netherlands, in which the Tegelen clay pits played
an important role. When BROUWER left the museum for a
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post at Leiden University, his position as curator was taken
over by KORTENBOUT VAN DER SLUIJS, who also participated
in the collecting trips. Apparently the more fossiliferous
beds mentioned by SCHREUDER were now being exploited.
Although mechanical digging had replaced the handwork
in many of the pits, the Leiden team succeeded in gathering the largest Tegelen collection. This should have been
a tremendous boost for the research, but things turned
out differently. VAN DER VLERK intended the study of the
Tegelen Clay, including its fossil remains, as a PhD thesis
for KORTENBOUT VAN DER SLUIJS (ZAGWIJN in litt. 2003). His
student, however, proved to be not much of a writer. He
did co-author an article on the stratigraphy of the Tegelen
Clay Pits (KORTENBOUT VAN DER SLUIJS & ZAGWIJN 1962),
and gave a description of the tapir fossils from Maalbeek
(KORTENBOUT VAN DER SLUIJS 1960), but other than that
produced no papers on the fauna. The research, however,
was claimed and would remain so until the beginning of
the 1980s.
At the beginning of the 1960s micromammal research
got a tremendous boost through the work of among others
VOLKERT FAHLBUSCH, PIERRE MEIN, HANS DE BRUIJN and
THIJS FREUDENTHAL. The latter was contacted by WALDO
ZAGWIJN of the Dutch Geological Survey. At the time the
former clay-pits near Tegelen were flooded, but ZAGWIJN
had received word that the pit Russel-Tiglia-Egypte would
be temporarily drained. During a visit to the pit TOM MEIJER,
a malacologist of the Geological Survey, had taken
samples from a filled-in stream gully in which he found a
number of rodent molars.
FREUDENTHAL started collecting at the Russel-TigliaEgypte pit in 1970. For seven years collecting continued
during campaigns, which lasted about three weeks each
(fig. 3). The goal of the campaigns was two-fold. Of
course, they were aimed at building a good collection of
Tegelen microvertebrates. FREUDENTHAL’s personal interest, however, lay with developing a method for processing
fossiliferous sediment, which he then could use in microvertebrate research on Miocene deposits in Spain.
FREUDENTHAL et al. (1976) published a preliminary
report on the field campaigns. In this article ALBERT VAN
DER MEULEN from Utrecht University presented the first
overview of the micromammal fauna. VAN DER MEULEN
had finished his PhD on Pleistocene faunas some years
earlier and therefore had a much better background for
working on the fauna than FREUDENTHAL, whose experiences lay with Miocene faunas. Thus, part of the material
from the campaigns was sent to Utrecht, both in sorted and
unsorted samples, whereas the rest was stored in Leiden.
Only in 2001 were all of the micromammals from Tegelen
registered. This involved not only the return of the Utrecht
collection to Leiden, but also the washing and sorting of
unprocessed sediments still in the museum. Only then did
the overall results of the campaigns become clear. FREUDENTHAL and his team had collected c. 5000 dental elements
of micromammals from the pit Russel-Tiglia-Egypte. In
order to do so, they had sieved c. 192 m3 of sediment in a
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VAN DEN
HOEK OSTENDE & DE VOS: A century of research on the classical locality of Tegelen
clay bed, which was about two metres high. On average,
24 micromammal fossils per m3 were collected, which suggests that the clay was poor in fossils. However, from the
quadrant system used by FREUDENTHAL, it can be calculated
that the majority of the molars come from the lowermost
part of the section, where the concentrations are well over
one hundred molars per m3. In contrast, some of the uppermost quadrants proved to be sterile.
The fauna
Fig. 3: In the 1970s nearly 200 m3 of clay from the pit RusselTiglia-Egypte was sieved in search of micromammals.
Non-mammalian vertebrates: Most of the papers on the
Tegelen fauna deal with the mammals, but these are
not the only vertebrates found in the Limburg locality.
GAUDANT (1979) distinguished eleven species of fish in
the Tegelen fauna. The assemblage consisted of species
still found in The Netherlands and was indicative for
slow moving to stagnant waters. The only reptile to be
described from Tegelen thus far is the European pond
turtle, Emys orbicularis, already noted by DUBOIS (1904).
SCHREUDER (1946) gave a description of a beautifully preserved carapace of this turtle. Preliminary identifications
made by MASSIMO DELFINO of Florence University show
that Tegelen had a rich herpetofauna, with at least thirteen
different species.
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Fig. 4: The first reconstruction of the Tegelen landscape as made by Dubois. Note the presence of a hippopotamus, based on the
false identification of Sus strozzii.
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Cour. Forsch.-Inst. Senckenberg, 256, 2006
The only bird described from Tegelen is Haliaaetus
albicilla. VAN REGTEREN ALTENA had found in the Tegelen
collection in Teylers museum several bones, which had
been found close together and presumably belonged to one
individual. The fossils were described by JUNGE (1953).
The Leiden collection harbours several unidentified bird
remains.
The large rodents: Remains of the extinct beaver Trogontherium cuvieri are so abundant in the Tegelen Clay that
in the German literature these beds are often referred to
as the “Trogontherium-Tone”. Apart from this beaver,
Castor fiber was also found at Tegelen, but it is far more
rare. The full description of the Tegelen beaver fossils was
given by SCHREUDER (1928, 1929, 1931). She used the name
Trogontherium boisvilletti, as she made a difference between a western and an eastern species of Trogontherium,
T. boisvilletti and T. cuvieri, respectively. MAYHEW (1978)
used the same names, but as chronosubspecies. Pre-Elsterian find (thus including Tegelen) were attributed to T.
c. boisvilletti, the younger representative to T. c. cuvieri.
HEINRICH (1998) did not make any distinctions between
subspecies, but showed the evolutionary developments
within T. cuvieri. His paper shows that the assemblage
from Tegelen is not only the most primitive one, but also
by far the oldest.
Another large rodent, Hystrix refossa, is represented
only by the articulated distal end of a humerus and proximal part of the ulna. BERNSEN (1932b) had described the
fossil as H. cf. etrusca, a junior synonym of H. refossa
(VAN WEERS 1994). The presence of a porcupine in Tegelen is remarkable. Present Hystrix favours dry and open
biotopes, whereas Tegelen represents a wet, forested
environment.
Primates: The presence of a monkey was first noted by
BERNSEN (1930a). Macaca sylvanus is a rare element in
the fauna, but is represented by one of the best preserved
fossils from Tegelen, a lower mandible with almost a
complete dentition. The mandible was preserved in a
concretion of clay, a mode of fossilisation not uncommon
in Tegelen. Originally BERNSEN described it as Macaca
florentina, but nowadays the Pleistocene monkeys of
Europe are considered to belong M. sylvanus (SZALAY &
DELSON 1979). SCHREUDER (1945, 1949) described some
additional material. A deciduous molar of the monkey was
found on the sieve during the micromammal expeditions
of the 1970s. Thus, the macaque is one of the few larger
mammals found in conjunction with the micromammal
fauna.
Artiodactyles: The deer were the first fossil mammals to
be described from Tegelen. Apart from Trogontherium
cuvieri, they are the most common large mammals in
the fauna. DUBOIS (1905) recognised three species. As
was usual in those days, small differences with fossils
from other localities were emphasised and Tegelen got
its ‘own’ species of deer, Cervus teguliensis and Cervus
rhenanus, the latter being named after the river Rhine,
which deposited the clay-beds. The third species present
eschweizerbartxxx sng-
according to DUBOIS was Cervus dicranius. The revision
of BERNSEN (1933/34), which was finished after his death
by SCHREUDER, made the situation even more complex,
since Cervus cf. ctenoides was added to the list. KUNST
(1937), however, assumed that only one large species
of deer was present in Tegelen, which she called Cervus
(Eucladoceros) teguliensis.
By the end of the 1980s it was clear that there was a
wild growth in the taxonomy of the Villafranchian deer.
The material from Tegelen was restudied, particularly in
comparison to the French Villafranchian species, which
had been extensively described by HEINTZ (1970). SPAAN
(1992) concluded that the French Cervus philisi was a
junior synonym of C. rhenanus, and that Eucladoceros
teguliensis and E. senezensis were also conspecific. Later,
a revision of the genus Eucladoceros showed that E. ctenoides was the oldest available name for this species (DE
VOS et al. 1995).
The first fauna lists from Tegelen (DUBOIS 1904, 1905)
featured Hippopotamus, and the animal also appears on
a reconstruction he made of the Tegelen environment
(fig. 4). The presumed presence of hippos in Tegelen was
based on the misidentification of a canine of the pig Sus
strozzii. Hippopotamus was also listed for Tegelen by
GUÉRIN (1980), but hippos have never been found in the
Tegelen Clay.
The presence of Sus strozzii was first noted by
RICHARZ (1921), who gave a short description of nine
molars. BERNSEN (1931c) described the finds in more detail. The best fossil of the Tegelen pig can be found in the
Amsterdam collection, to which it was donated in 1935.
It is an almost complete mandible of a young individual
and was described by SCHREUDER (1936a, 1945).
Leptobos is a rare element in the Tegelen fauna.
SCHREUDER (1936a) described some molars, a horn pit
and a scapula. HOOIJER (1947) described a radius and
KORTENBOUT VAN DER SLUIJS (1960) mentioned the presence
in the Maalbeek pit. The finds from Tegelen are attributed
to Leptobos elatus.
Perissodactyles: BERNSEN (1927) wrote a PhD on the
rhinoceroses of the Tegelen Clay. He concluded that
there were two species of rhino represented in the fauna,
viz. ‘Rhinoceros’ etruscus and ‘Rhinoceros’ Mercki. It is
noteworthy that all of the material assigned to the second
species by BERNSEN came from the same collection, that
from the Mission Museum at Steyl (now included in the
Naturalis collection). Some of the rhinoceros remains in
the Natuurhistorisch Museum Maastricht were identified
by SCHREUDER as ‘merkii’. LOOSE (1975) did not concur
with BERNSEN, and believed that all of the material should
be assigned to a single species, Dicerorhinus etruscus. But
GUÉRIN (1980) also recognised the two species found by
BERNSEN, nowadays known as Stephonorhinus etruscus
and S. kirchbergensis.
So it is a matter of debate whether or not two species
of rhinoceros are represented. This is partly due to the
nature of the material, which is fragmented and not very
297
VAN DEN
HOEK OSTENDE & DE VOS: A century of research on the classical locality of Tegelen
rich. Therefore, it seems open to different interpretations.
On the other hand, one would not expect Stephanorhinus kirchbergensis, a Middle Pleistocene species, in the
Tegelen fauna. As GUÉRIN interpreted the fauna to be
much younger than is commonly assumed, this was not as
much a problem for him. However, this interpretation of
the age was partly based on the false assumption that the
hippopotamus was present in the Tegelen fauna.
Horse fossils are rare in Tegelen, but the presence of
equids was already noted by DUBOIS (1904). He classified
the horse as E. stenonis COCCHI race major BOULE. The
same classification was used by BERNSEN (1931a). When
RAVEN (1935) described large metapodials from Tegelen,
he preferred the name Equus robustus. The use of different
names is indicative of the confusion in horse taxonomy at
the time. VIRET (1954) introduced the name E. bressanus,
with which he specifically indicated not only the finds from
Chagny and Sénèze, but also the remains from Tegelen.
In 1930 some peculiar molars were found near Maalbeek by A. BLOEMERS, a clay worker for the N.V. Nederlandse Gresbuizenindustrie. He took the fossils home,
where they lay for several decades. In the 1950s, when
he was nearly 90 years old, he donated the fossils to a
psychologist in Belfeld, Dr. VERKOOIJEN, for whom he was
a gardener. VERKOOIJEN took the fossils to the Natuurhistorisch Museum Maastricht for identification, where the
curator E. M. KRUYTZER identified them as the dentition of
a tapir. The exact locality where the molars had been found
is not clear. BLOEMERS was asked to indicate the place he
had excavated the molars. Unfortunately, the account given
by KRUYTZER (1957) in his preliminary note differs from
the one given by KORTENBOUT VAN DER SLUIJS (1960), who
gave a full description of the fossils. Both authors agree,
however, that the find was made in the top layer of a blue
clay, just below a brown bank.
eschweizerbartxxx sng-
Carnivores: The carnivore guild of Tegelen consists of
one bear, one hyena, one cat and two mustelids. No species
of dog were found and other unexpected absentees are the
sabre-toothed cats. The bear Ursus etruscus was the first
carnivore to be described (NEWTON 1913). BERNSEN (1932a)
also described canines from the bear from the collection
of the Natuurhistorisch Museum Maastricht. SCHREUDER
(1935) described a second dentition of U. etruscus, and
gave a description of a third dentition in 1949. That year,
two mandibles were found. It is clear from notes in the registration books in Naturalis that SCHREUDER had seen these
specimens, but they were never described. No post-cranial
elements of the bear have been collected at Tegelen.
The hyena is represented both by coprolites and skeletal elements. The first description of the hyenid fossils
was given by BERNSEN (1931d). SCHREUDER (1949) noted a
peculiarity about the hyena fossils from Tegelen. There is
a large variation in the sizes and, particularly, a dentition
in the Maastricht museum stands out by being very small.
SCHREUDER explained this by assuming sexual dimorphism.
This provides, however, a problem, since recent hyenas
298
show only a limited size difference between males and
females. According to HOLEKAMP & SMALE (2000) females
of Crocuta crocuta are only 10% larger than the males, a
difference which would be unnoticeable in fossil assemblages. BERNSEN (1931d) and SCHREUDER (1949) classified
the species as Hyaena perrieri. WERDELIN & SOULANIAS
(1991) listed the Tegelen assemblage under the name
Pliocrocuta perrieri.
SCHREUDER (1935) indicated the presence of two
mustelid species in Tegelen. However, the material she
described all belongs to a single species, Enhydrictis
ardea. Nevertheless, a second mustelid is present. WILLEMSEN (1988) described, apart from the Enhydrictis fossils, a mandible of the pole-cat Mustela palerminea. This
mandible had been obtained during one of the collecting
trips of the curators from the Leiden Museum and is now
in the collection of Naturalis.
The pantherine fossils from Tegelen were described
for the first time by VON KOENIGSWALD (1961), who named
the panther Panthera schreuderi (sic). HEMMER & SCHÜTT
(1969) synonymized Panthera schreuderi with P. gombaszoegensis. However, according to HEMMER (1965), one
of the specimens from Tegelen represented P. schaubi, a
rather enigmatic species from Séneze, which was later
synonymised with Puma pardoides (HEMMER 2001). HEMMER et al. (2004) explicitly included part of the Tegelen
material in this species.
O’REGAN & TURNER (2004) gave full descriptions of all
available specimens, including a cast made from a natural
mould formed by a concretion. They concluded that all of
the material could be attributed to one species. However,
like in the hyenas, there is a remarkable variation in size.
O’REGAN & TURNER provided several possible explanations
for this, including the possibility that the material represented different time frames (P. gombaszoegensis shows
an increase in size over time). In the end, they decided the
best explanation for the differences in size was assuming
sexual dimorphism.
Proboscideans: Remains of elephants from the Tegelen
Clay are rare. Only eleven fossils have been found, all
of which can be attributed to Mammuthus meridionalis
(GUENTHER 1986). In what is generally referred to as ‘the
main fauna’, that is the finds from the direct vicinity of
Tegelen, only one proboscidean species has been found.
However, in 1960 a molar of the mastodont Ananacus
arvernensis was found in the Maalbeek pit, somewhat
more to the south. Originally the find was dated as Eburonian, based on the pollen analysis of the clay adhered to the
molar (KORTENBOUT VAN DER SLUIJS & ZAGWIJN 1962). The
pollen spectrum from this clay clearly represented a cold
period. WESTERHOFF et al. (1998), however, showed that
the find layer was probably deposited in a colder period
within the Tiglian, older than the main fauna.
M i c r o m a m m a l s : The presence of micromammals in
the Tegelen Clay was first noted by NEWTON (1907), who
described finds of the vole Mimomys pliocaenicus.
Cour. Forsch.-Inst. Senckenberg, 256, 2006
As we noted earlier, no palaeontological excavations
were organised at Tegelen and thus for over half a century
nobody took to sieving the clay in search of micromammals. Nevertheless, chance finds did occur, and ANTJE
SCHREUDER (1945: 18) praised the clay workers that had
found fossils of hares and desmans. She extensively
discussed the find of the hare Hypolagus brachygnathus
(SCHREUDER 1936b) and described the desmans from
Tegelen in her review of this subfamily of Talpidae
(SCHREUDER 1940).
Systematic study of the micromammals only became
possible after the expeditions of FREUDENTHAL in the 1970s.
The first large paper dealing with small mammals from the
Limburg locality was the PhD thesis of REUMER (1984),
in which he compared the Tegelen shrews to those from
various Hungarian localities. REUMER noted the presence
of four shrew species: Beremendia fissidens, Petenyia hungarica, Sorex minutus and Drepanosorex praearaneaus.
The latter species is nowadays generally placed in Sorex,
of which Drepanosorex is considered a subgenus (REUMER
1985). RÜMKE (1985) included the desmans from Tegelen
in her review on the Neogene Desmaninae. Two desman
species are present in the fauna, Desmana thermalis and
Galemys kormosi. A student of RÜMKE, JOLANDA RODERS,
wrote an unpublished MSc thesis on the talpine mole from
the locality and concluded that the assemblage is referable
to Talpa minor (RODERS 1987).
The vole fauna was studied by TESAKOV (1998). The
voles are represented by six species, Mimomys pliocaenicus, M. tigliensis, M. reidi, Pitymimomys pitymyoides,
Clethrionomys kretzoii and Ungaromys nanus. The studies
of REUMER, RÜMKE and TESAKOV were all done before all of
the material had been sorted. Since then, the material has
nearly been doubled. Among the material found there were
five molars of the lemming Dicrostonyx. Unfortunately,
it is unclear from which quadrant these fossils came. The
find of Dicrostonyx in a fauna otherwise dominated by
elements typical for a forest environment is surprising,
and as yet not understood.
The recently screened sediment harboured another surprise in the glirids, viz. the first record for The Netherlands
of Glirulus pusillus. This small dormouse was, together
with the far more numerous Muscardinus pliocaenicus,
described by VAN DEN HOEK OSTENDE (2003). The sciurid
from Tegelen presented even a bigger surprise; the species
identified as Sciurus cf. vulgaris by FREUDENTHAL et al.
(1976) turned out to be a new, large species of Hylopetes,
H. debruijni (REUMER & VAN DEN HOEK OSTENDE 2003).
FREUDENTHAL et al. (1976) already noted the presence of
the murids Apodemus and Micromys in Tegelen. Apodemus
is the most numerous micromammal in the Tegelen fauna.
The bats from Tegelen are as yet undescribed.
Micromammals were also collected from the pit Maalbeek (WESTERHOFF et al. 1998). This assemblage is older
than the one collected from the pit Russel-Tiglia-Egypte,
but contains the same species. The studies in the Maalbeek pit are still continuing under the direction of THIJS
eschweizerbartxxx sng-
VAN KOLFSCHOTEN of the Archaeological Institute of the
Leiden University.
Table 1 gives a full overview of the Tegelen fauna. The
finds from Maalbeek have not been included in this table,
since it has been clearly established that they are older
than the main fauna (WESTERHOFF et al. 1998).
The age of the Tegelen fauna
The age of the Tegelen fauna seems easily enough determined. It is, of course, Tiglian. However, that answer
is a bit too easy. We first have to determine that there is
indeed one Tegelen fauna and secondly, since the Tiglian
and its boundaries are not defined by fossil mammals, the
stage as such cannot be used for interregional correlations
in mammal palaeontology.
As to the question whether or not there is one Tegelen
fauna, it is obvious that not all of the finds from the Tegelen area are of the same age. The fossils of the tapir and
mastodont f ound in the pit Maalbeek are generally considered to be older than the fauna found in the pits nearer
to the village of Tegelen itself (WESTERHOFF et al. 1998).
Of course, these rare species attract attention, but they are
certainly not the only large mammals found in Maalbeek.
KORTENBOUT VAN DER SLUIJS (1960) also mentioned the
presence in Maalbeek of Eucladoceros ctenoides, Cervus
rhenanus, Leptobos cf. elatus and Stephanorhinus etruscus, species that are also known from other Tegelen pits.
Another obvious difference in age lies with the micromammals. These were collected from the filled in stream
gully in the pit Russel-Tiglia-Egypte and are thus per
definition separated from the main clay beds in that pit.
However, the hiatus between the deposition of the two
is of unknown duration. The Tiglian substages TC5 and
TC6 were defined on the basis of pollen spectra in the
stream gully from which the micromammals were collected. Most of the material was, together with a rich seed
flora, collected from the base of this gully. The recognised
evolutionary stages of all recorded arvicolid species and
the lack of remains of Microtus place the fauna in the
Late Villanyian (TESAKOV 1998). This age is corroborated by, for example, the stage of evolution of the glirid
Muscardinus pliocaenicus (VAN DEN HOEK OSTENDE 2004).
However, its relation to the larger mammal fauna from the
region is unclear. Furthermore, after TESAKOV published
his findings on the voles from Tegelen, five molars of the
lemming Dicrostonyx were found in the collection. This
suggests that even in the small mammal assemblage some
contamination may be present, since a lemming does not
seem to fit the remainder of the micromammal assemblage,
which consists primarily of wood-inhabitants and/or semiaquatic species.
Apart from the older age for the fauna from the
Maalbeek pit, and the (somewhat) younger age of the
micromammal fauna, the homogeneity of the other fossils is also open to doubt. The reason for this lies with the
299
VAN DEN
HOEK OSTENDE & DE VOS: A century of research on the classical locality of Tegelen
Table 1: Faunal list of Tegelen. The second column gives the last description. If the classification used in this paper deviates from
the one in the last description, the reference to the classification is added in the third column. Species exclusively found in the
Maalbeek pit (Anancus arvernensis, Tapirus arvernensis) have not bee included in the list.
Description
Classification
according to
Description
Hylopetes debruijni
REUMER &
VAN DEN HOEK
OSTENDE 2003
Muscardinus pliocaenicus
VAN DEN HOEK
OSTENDE 2003
Glirulus pusillus
VAN DEN HOEK
OSTENDE 2003
GAUDANT 1979
Micromys sp.
FREUDENTHAL et
al. 1976
Scardinus erythophthalmus
GAUDANT 1979
Apodemus cf. sylvaticus
Rutilus rutilus
GAUDANT 1979
FREUDENTHAL et
al. 1976
Mimomys pliocaenicus
TESAKOV 1998
Mimomys tigliensis
TESAKOV 1998
Mimomys reidi
TESAKOV 1998
Pitymimomys pitymyoides
TESAKOV 1998
Clethrionomys kretzoii
TESAKOV 1998
Ungaromys nanus
TESAKOV 1998
Dicrostonyx sp.
Unpublished
Hystrix refossa
BERNSEN, 1932b
PISCES
Esox lucius
GAUDANT 1979
Anguilla anguilla
GAUDANT 1979
Carassius carassius
GAUDANT 1979
Tinca tinca
GAUDANT 1979
Abramis brama
GAUDANT 1979
Alburnus alburnus
Cyprinidae gen. et sp. indet. GAUDANT 1979
Perca fluviatilis
GAUDANT 1979
Lucioperca lucioperca
GAUDANT 1979
Gasterosteus aculeatus
GAUDANT 1979
AMPHIBIA
Triturus cristatus
det. M. DELFINO
Triturus sp.
det. M. DELFINO
Palaeobatrachidae indet.
det. M. DELFINO
Lagomorpha
Bufonidae indet.
det. M. DELFINO
Hypolagus brachygnathus
Rana gr. R. ridibunda
det. M. DELFINO
Hyla gr. H. arborea
det. M. DELFINO
Pelobates sp.
det. M. DELFINO
Bombina sp.
det. M. DELFINO
VAN WEERS 1994
SCHREUDER 1936b
Chiroptera
eschweizerbartxxx sng-
Chiroptera undet. 1
Unpublished
Chiroptera undet. 2
Unpublished
Primates
Macaca sylvana
REPTILIA
Emys orbicularis
SCHREUDER 1945
Anguis fragilis
det. M. DELFINO
Mammuthus meridionalis
Lacerta sp.
det. M. DELFINO
Artiodactyla
Colubrines indet.
det. M. DELFINO
Sus strozii
Natrix natrix
det. M. DELFINO
Eucladoceros ctenoides
SCHREUDER 1945,
1949
SZALAY & DELSON
1979
Proboscidea
AVES
Haliaaetus albicilla
Classification
according to
JUNGE 1953
GUENTHER 1986
SPAAN 1992
Cervus rhenanus
SPAAN 1992
Leptobos cf. elatus
SCHEUDER 1936a,
HOOIJER 1947
DE VOS et al. 1995
Perissodactyla
MAMMALIA
Stephanorhinus etruscus
GUÉRIN 1980
Soricomorpha
?Stephanorhinus
kirchbergensis
GUÉRIN 1980
BERNSEN 1931a,
RAVEN 1935
Desmana thermalis
RÜMKE 1985
Galemys kormosi
RÜMKE 1985
Equus robustus
Carnivora
Talpa minor
RODERS 1987
Petenyia hungarica
REUMER 1984
Mustela palerminea
WILLEMSEN 1988
Beremendia fissidens
REUMER 1984
Enhydrictis ardea
WILLEMSEN 1988
Sorex (Drepanosorex)
praearaneus
REUMER 1984
Ursus etruscus
SCHREUDER 1945,
1949
Sorex minutus
REUMER 1984
Panthera gombaszoegensis
O‘REAGAN &
TURNER 2004
Pliocrocuta perrieri
SCHREUDER 1949
REUMER 1985
VIRET 1954
Rodentia
Trogontherium cuvieri
SCHREUDER 1928
Castor fiber
SCHREUDER 1928
300
MAYHEW 1978
WERDELIN &
SOULANIAS 1991
Cour. Forsch.-Inst. Senckenberg, 256, 2006
carnivores and the rhinoceroses. In the case of the rhinos, it
is the assumed presence of Stephanorhinus kirchbergensis
(BERNSEN 1927, GUÉRIN 1980) that presents a problem. This
species is otherwise known from the Middle Pleistocene
and Tegelen would be by far the oldest occurrence. Such
a very early occurrence seems unlikely. One possible explanation would be that a part of the fossils from Tegelen
is indeed much younger than the rest. In favour of this
hypothesis would be that almost all of the S. kirchbergensis fossils are from one collection (viz. Mission Museum
Steijl), and thus likely to have been collected at the same
locality. The alternative explanation would be that the presence of S. kirchbergensis is based on the misidentification
of S. etruscus fossils, as assumed by LOOSE (1975).
The large carnivores from Tegelen all show a large
variation in size. SCHREUDER (1949) explained this variation
in the bears and hyenas by assuming sexual dimorphism,
and the same explanation was coined for the panther by
O’REGAN & TURNER (2004). In the case of the bears, sexual
dimorphism is well documented. However, for hyenas the
very limited dimorphism found in recent forms makes it
an unlikely explanation for the large variation found in the
small fossil assemblage. In our opinion, the same holds
true for the panther. O’REGAN & TURNER (2004) found the
amount of variation acceptable, since it is only somewhat
larger than the differences between males and females
in lions. However, the sample they compared with was
much larger than the few fossils available from Tegelen.
Furthermore, the lion may not be the best felid to compare
with, since it is the only pack hunter in its family, with
different roles for males and females. O’REGAN & TURNER
(2004) did consider the possibility of different ages in the
Tegelen fauna, since Panthera gombaszoegensis shows an
increase over time. However, they rejected this on the basis
of Tegelen being a classical locality. Given the uncertainty
about the provenance of the material, we feel that assuming
a different age for various parts of the fauna is at least as
good an explanation as sexual dimorphism.
Another peculiarity about the Tegelen fauna is the
presence of Trogontherium cuvieri. This extinct beaver
has its first occurrence in the Limburg locality and the
assemblage is also clearly the most primitive one (HEINRICH 1998). The age difference with the next occurrence
discussed by HEINRICH is, however, remarkably large. The
assemblage from the Maasvlakte is about one million years
younger than the assumed age of Tegelen. In other words,
like Stephanorhinus kirchbergensis, T. cuvieri is a species
otherwise known from the Middle Pleistocene.
Thus, there are several indications that the fossils from
Tegelen come from different time frames. The evidence
is somewhat circumstantial, but this is logical given the
relatively stability of the Villafranchian ecosystem. If for
example, the pit Maalbeek had not yielded remains of
mastodont and tapir, the remainder of the fauna would
not have given any reason to assume an older age of
the sediments in that particular pit. By the same token,
younger sediments in the area would yield a very similar
eschweizerbartxxx sng-
fauna. Only evolutionary changes within lineages, such as
the increase in size of various carnivores, would reveal a
difference in age.
To complicate things further, the mammal sequence
in the Netherlands is very incomplete. There appears to
be a hole of one million years between Tegelen and the
next faunas, which are assigned to the Bavelian, viz. the
Maasvlakte (VAN KOLFSCHOTEN & VERVOORT-KERKHOFF
1999) and Het Gat (MOL et al. 2003). These faunas are not
even in situ, but have been found washed up on the beach
(Maasvlakte) or have been collected by fishermen from the
bottom of the North Sea (Het Gat). There are, however,
some Bavelian fossils that have been found in situ in The
Netherlands. Two different locations near Bavel yielded
a small mammal assemblage and some macrofossils (VAN
KOLFSCHOTEN 1990). Even in the Zuurland boreholes this
gap of a million years seems present, as the ‘Tiglian’
faunas are immediately followed by ‘Cromerian’ faunas
(REUMER & HORDIJK 1999).
The presumed one million year gap is relevant for the
interpretation of the Tegelen fauna. So far the fauna has
always been interpreted as a single fauna, its upper limit set
by the upper boundary of the Tiglian. Thus, the possibility
that part of the material was derived from younger sediments in the upper part of the Waalre formation was never
taken into account. Such an explanation would explain the
peculiarities in the mammal record. This would imply that
in fact the one million year gap has been partly filled, but
that this has never been recognised. Given the stability
of the Villafranchian mammal community over time, we
would even not expect to be able to recognise ‘Tiglian’
mammal fauna from younger ones. The main argument
against the presence of younger sediments is provided
by palaeomagnetism. VAN MONTFRANS (1971) found a
normal polarity in various pits in which mammal fossils
have been found. DREES (2005) noted that the presumed
reversal to reversed polarity on the Tiglian-Eburonian
boundary is based on sequences, the age of which had
been reinterpreted since the publication of MONTFRANS’
thesis. Nevertheless, it is most likely that the sediments
in the pits Egypte, Kurstjens, Wambach, and Laumans
were deposited during the Olduvai Chron (Chron C2n),
providing an upper age limit of 1.77 My for the mammals
from these pits.
Conclusions
A century of research on the clay-pits near Tegelen has
yielded extensive collections of Villafranchian mammals
in various Dutch museums (Teylers Museum, Naturalis,
Zoological Museum Amsterdam, Natuurhistorisch Museum Maastricht). We owe these collections to the ceramic
industry in the region, as clay workers regularly gathered
chance finds of bones and teeth. No scientific excavations
have ever taken place and, therefore, the exact provenance
of the material is not known (although usually the pit
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VAN DEN
HOEK OSTENDE & DE VOS: A century of research on the classical locality of Tegelen
where it has been collected is indicated). Comparing the
number of fossils retrieved to the huge amount of sediment
excavated during the 20th century suggests that the clay
beds near Tegelen may not have been very rich in fossils.
From notes by SCHREUDER we know that some of the clay
beds had a much richer fossil contents than others. Her
writings also provide an explanation for the fragmentary
nature of most of the Tegelen fossils. The bones were
often waterlogged and as they were pulled out of the clay
by laymen, they were often damaged during excavation.
Therefore it is certainly no coincidence that some of the
finest fossils from Tegelen were encrusted in concretions,
protecting them from harm by injudicious handling by
clay-workers.
The only scientific excavations that did take place
near Tegelen were the expeditions in search of smaller
mammals in the pit Russel-Tiglia-Egypte (FREUDENTHAL et
al. 1976) and the Maalbeek pit (WESTERHOFF et al. 1998).
The assemblage from Russel-Tiglia-Egypte was sampled
from a 2 m clay bed in a filled-in stream gully. Most of
the assemblage springs from the bottom of this in-fill.
The age of the assemblage was determined as Upper Villanyian on the basis of the state of evolution of Mimomys
pliocaenicus (TESAKOV 1998). The recent discovery of five
Dicrostonyx molars in the assemblage is a puzzle that is
as yet unsolved.
As we do not know the exact location in which the
various large mammal fossils were found, the question
whether or not they are all of similar age is justified. The
presumed presence of Stephanorhinus kirchbergensis
(GUÉRIN 1980), and interpreting the large variations in size
within the panther and hyena as a state of evolution rather
than the result of sexual dimorphism, would suggest that
indeed material of different ages is mixed. Although the
evidence of mixture of ages is circumstantial, we suggest
that there are enough doubts to be cautious about the age
of Tegelen, when using it for interregional correlations.
eschweizerbartxxx sng-
Acknowledgements
Various people helped us with our research and with finishing our manuscript. ANNE SCHULP provided us with a copy
of the registration data on Tegelen of the Natuurhistorisch
Museum Maastricht. MASSIMO DELFINO kindly allowed
us to use his identifications of the herpetofauna. WILLEM
KAKEBEEKE reconstructed the micromammal excavation
from FREUDENTHAL’s notes, and calculated the fossil richness of the various quadrants. TOM MEIJER critically read the
manuscript, and provided us with some useful comments.
DORIS NAGEL suggested and provided copies of literature on
hyenas, and helped formulate the German abstract. STEVE
DONOVAN made some linguistic improvements; additional
linguistic remarks were made by PAUL MAZZA. RALF-DIETRICH KAHLKE and LUTZ MAUL acted as reviewers, and suggested a number of improvements to the text. The help of
all these colleagues was greatly appreciated.
302
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Manuscript submitted 2005–10–04
Revised manuscript accepted 2006–06–12