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Survival of the thinnest: rediscovery of Bauer’s (1898) ichthyosaurtooth sections from Upper Jurassic lithographic limestonequarries, south Germany
Torsten M. Scheyer • Markus Moser
Received: 12 July 2011 / Accepted: 12 September 2011
� Swiss Geological Society 2011
Abstract The re-discovery of nine petrographic slides
from the late 19th century at the palaeontological collec-
tions of the University of Zurich, showing thin-sectioned
ichthyosaur teeth, revealed these slides be the only pre-
served remains of the historical collection of Upper
Jurassic ichthyosaurs from the Bavarian State Collection
for Palaeontology and Geology; fossil material which, up
to now, was thought to have been completely destroyed
during World War II. Here the history of these slides, from
their origin in Munich as part of the doctoral thesis of Franz
Bauer (1898) to their rediscovery in Zurich in 2010 is
presented. Furthermore, a complete overview of all slides
is given to elucidate their scientific value with the back-
ground of up-to-date knowledge of ichthyosaur dentition
and tooth histology, including aspects of tissue and growth
mark identification. As such, the sectioned teeth show an
exposed layer of acellular cementum at the tooth neck, and
sets of short and long period growth lines in the ortho-
dentine. The slides of one tooth are part of the original
syntype material of Aegirosaurus leptospondylus (WAGNER).
They reveal an oval rather than a rectangular shape of the
root, as well as the presence of peculiar vascular canals,
interpreted as secondary osteodentine deposition, in the
peri-pulpal orthodentine.
Keywords History of science � Ichthyopterygia �Ichthyosaurus trigonus var. posthumus �Aegirosaurus leptospondylus � Nannopterygius �Orthodentine � Growth increments
Institutional abbreviations
BSPG Bavarian State Collection for Palaeontology
and Geology, Munich = Bayerische
Staatssammlung fur Palaontologie und
Geologie, Munchen, Formerly Bayerische
Palaontologische Staatssammlung
PIMUZ Palaontologisches Institut und Museum,
University of Zurich, Switzerland
Introduction
General introduction
In 2010, a tray with nine petrographic thin-sections of two
ichthyosaur teeth was discovered during renovation works
in the Palaeontological Institute and Museum of the Uni-
versity of Zurich. From the labels still attached to the slides
it was apparent that these slides were part of the doctoral
thesis materials of Dr. phil. Franz Bauer in Munich, pub-
lished in Palaeontographica in 1898. Besides the nine
slides still preserved (Fig. 1), all other remains of the
ichthyosaur specimens figured on plates 25–27 by Bauer
(1898) were completely destroyed during World War II
(WWII) on the 24–25. April 1944 during a bombing raid
of the old Academy building in Munich, which housed
the ‘Bayerische Palaontologische Staatssammlung’ (now
Editorial handling: Michael J. Benton & Daniel Marty.
Electronic supplementary material The online version of thisarticle (doi:10.1007/s00015-011-0076-y) contains supplementarymaterial, which is available to authorized users.
T. M. Scheyer (&)
Palaontologisches Institut und Museum der Universitat Zurich,
Karl Schmid-Strasse 4, 8006 Zurich, Switzerland
e-mail: [email protected]
M. Moser
Bayerische Staatssammlung fur Palaontologie und Geologie,
Richard-Wagner-Straße 10, 80333 Munich, Germany
Swiss J Geosci
DOI 10.1007/s00015-011-0076-y
BSPG) at that time. The specimens were stored and dis-
played in public exhibition then (Dehm 1978). Similarly,
all paper documents, including inventory, loan documents,
field books, working notes, manuscripts, and theses, were
lost to fire.
Bauer’s monograph (1898) work was often cited in the
palaeontological literature (e.g. Bauer 1900b; Broili 1907;
von Huene 1922; Kuhn 1934; Camp 1942). Especially
references to the osteological description of the caudal
region and tail fin referred to Ichthyosaurus trigonus var.
posthumus by Bauer (1898) already appeared shortly after
the original print in a review by Koken (1901, p. 476), the
monographic work on Triassic ichthyosaurs by Merriam
(1908, p. 40), and in palaeobiological textbooks (e.g. Abel
1919, pp. 461, 463). The histological sections of the teeth
shown in this work, however, went without further notice
for some decades. Peyer (1945) was the first to mention and
reproduce part of the histological details, i.e. figure 28 of
plate 26, of Bauer (1898), when discussing the presence of
minute bore canaliculi of Mycelites ossifragus Roux, 1887
(filamentous fungi = ‘‘kalkfressende Algen’’ of Bauer) in
ichthyosaur teeth and other vertebrate hard tissues. In the
same context, the figure was also reproduced in Peyer’s
(1968) book on comparative odontology. Bauer (1898,
p. 290) already interpreted the bore canaliculi as secondary
disease symptoms instead of primary or post-mortem
structures, a view which was validated by microstructural
and ultrastructural studies of fungoid lesions in extant fish
teeth (Schmidt 1954; Kerebel et al. 1979).
In 1947, Albert Besmer, published a doctoral thesis on
the dentition of ichthyosaurs under the supervision of Prof.
Bernhard Peyer. For this work, Besmer borrowed the ori-
ginal slides of Bauer (1898) for comparison to his material,
which was composed of ichthyosaurs from the Middle
Triassic of Monte San Giorgio, Ticino, Switzerland and
from the Lower Jurassic (Lias e) of Holzmaden in Baden-
Wurttemberg, southwestern Germany. Although the slides
had to be sent to Besmer prior to April 1944, it is unclear in
which year exactly the slides were sent to Zurich, because
there are no written loan forms or receipts from that time.
Besmer, however, in his thesis acknowledged the late Prof.
Broili, who apparently was responsible for the loan. Broili
retired in 1939 but he was working as a volunteer for the
State Collection until 1942, so probably the slides were
sent to Zurich before that time. Besmer noted the pro-
longed time it took to complete his thesis due to obstacles
caused by WWII (Besmer 1947, preface on p. 2, p. 7).
After 1947, the slides remained first in the ‘‘Zoologisch-
vergleichend anatomische Institut’’ and were stored and
finally ‘‘forgotten’’ for seven decades in an unnamed
drawer of the PIMUZ, which was formally established
under that name in 1956.
Besides the two passages mentioned above, Besmer
(1947) included two more references to the slides, namely
on page 3 (Bauer 1898, plate 26, figs. 28–30) when dis-
cussing how Bauer (1898) already identified Kiprijanoff’s
(1881) misidentification of bore canaliculi of Mycelites
ossifragus as larger dentine tubules, as well as on page 11
(Bauer 1898, plate 26, figs. 28–30) when noting the overall
similarity of his tooth sections to previously published
accounts by Bauer (1898), Fraas (1891), Kiprijanoff (1881)
and Owen (1840–1845). The osteological descriptions of
Bauer (1898) are furthermore referenced in Bardet and
Fernandez (2000) and McGowan and Motani (2003), when
discussing the assignment of the species studied by Bauer
Fig. 1 Photographs of all recovered slides of Bauer (1898). a1–a8BSPG AS XIX 504a–h (slides 1–7 and ‘‘8’’), tooth from the
‘‘Oberndorfer specimen’’, syntype material of Aegirosaurus lepto-spondylus (WAGNER, 1853) in Wagner (1853a), sectioned transversely
and longitudinally. b BSPG AS I 1656, longitudinal section of a tooth
from the ‘‘Solnhofen specimen’’, referable to Ichthyosauria indet., aff.
Nannopterygius
T. M. Scheyer, M. Moser
(see ‘‘Materials and methods’’ below), but the thin-sections
were not, for the reasons described above. Further mention
of Aegirosaurus leptospondylus (WAGNER) material either
in faunal or systematic overviews or in comparison to other
material is found in Kuhn (1957, 1961, 1968, 1971),
McGowan (1976), Buchy and Lopez Oliva (2009), Maisch
(2010), Angst et al. (2010), Fischer (2011) and Fischer
et al. (2011). The ichthyosaur material described and fig-
ured by Bauer (1898) as ‘‘Ichthyosaurus trigonus Owen
var. posthumus Wagner’’ belongs, according to Bardet and
Fernandez (2000), to three different specimens, which
represent three separate taxa: (1) the ophthalmosaurid
Aegirosaurus leptospondylus (the Oberndorfer specimen),
(2) Ichthyosauria indet. close to Caypullisaurus or Oph-
thalmosaurus (the Haberlein specimen), (3) Ichthyosauria
indet. aff. Nannopterygius (the Solnhofen specimen).
Concerning the Oberndorfer specimen, Bardet and
Fernandez (2000, p. 510) noted that the ‘‘specimen roughly
shares the same character combination that the two new
skeletons [the specimen in private col. Schwegler, desig-
nated as ‘‘neotype’’ and referred to as ‘‘SM unnumbered’’;
and BSPG 1954 I 608, designated as referred specimen by
Bardet and Fernandez] here described. The specific name
leptospondylus is thus kept as valid and newly combined to
Aegirosaurus […].’’ Concerning the Solnhofen specimen,
they further write that ‘‘because of the scarcity of the
illustrations given by Bauer (1898) it is not possible to
assign the Solnhofen specimen with certainty to this genus
[to Nannopterygius] and it is thus only considered as an
indeterminate ichthyosaur, possibly close to Nannoptery-
gius.’’ (Bardet and Fernandez 2000, p. 511).
According to McGowan and Motani (2003, p. 118),
Aegirosaurus leptospondylus (WAGNER, 1853), originally
described in Wagner (1853a) and as described and newly
combined by Bardet and Fernandez (2000, p. 504), is still
valid, and both Aegirosaurus BARDET AND FERNANDEZ, 2000
from the Solnhofen Formation, Upper Jurassic (Early
Tithonian), Bavaria, Germany, and Nannopterygius von
Huene 1922 from the Kimmeridge Clay; Upper Jurassic
(Kimmeridgian), Dorset, UK, belong to Ophthalmosauridae
BAUR, 1887.
After 70 years of storage in Zurich (PIMUZ), all slides
are again stored in the Bavarian State Collection for Pal-
aeontology and Geology, Munich (BSPG).
Material of the Haberlein specimen, the oldest finding of
an Upper Jurassic ichthyosaur, which was already described
by Quenstedt (1851–1852), was not sectioned histologically
by Bauer (1898), and is not further considered here.
The importance of the rediscovery and notification of
some of the oldest, still preserved sections of ichthyosaur
teeth from the 19th century, being part of the doctoral
thesis documentation on one side and being the sole
‘‘survivors’’ of the original specimens of Upper Jurassic
ichthyosaurs from the Bavarian lithographic limestone
quarries stored in Munich on the other, is obvious. Given
the fact that eight of these slides belong to the purportedly
lost original type material of Ichthyosaurus leptospondylus
WAGNER, 1853 (Wagner 1853a, b), a note on the taxonomic
status of the material is warranted and is in preparation
elsewhere. Besides reconstructing the history of the slides,
the purpose of this contribution is to give a complete
overview of all slides preserved, and to elucidate the value
of the slides with the background of up-to-date knowledge
of ichthyosaur dentition and tooth histology, including
aspects of tissue and growth mark identification.
Biographical data on Bauer and Besmer
Franz Bauer
Franz Bauer (see Electronic Supplementary Material for a
more exhaustive biography and bibliography of Franz
Bauer) was born in Dollnstein near Solnhofen, studied
theology in Eichstatt and worked there as a priest in pastoral
care before studying natural sciences in Munich in 1895. He
completed a doctoral thesis under Prof. Karl Alfred von
Zittel in 1897. From 1901, Bauer had an appointment as
associate professor of Geology and Palaeontology at the
Chemistry Department, Technical University of Munich.
He published several scientific articles and abstracts,
including anthropological themes (Bauer 1900a) and three
on palaeontological issues (Bauer 1898, 1900b, 1901 [the
latter being his postdoctoral/habilitation thesis]), before
having a tragic accident in 1903 on the Risserkogel
Mountain near Tegernsee, south of Munich.
Albert Besmer
Albert Besmer (1900–1981) was born in Zurich, Switzer-
land, studied at the Universities of Zurich and Fribourg,
and since 1928 worked as a dentist in Zurich. He finished
his doctoral thesis under the supervision of Prof.
Dr. Bernhard Peyer (Zoological Institute) at the Medical
Faculty of the University of Zurich in 1947. To our
knowledge, Besmer published only his thesis, for which the
original slides from Bauer (1898) were sent to him by Prof.
Dr. Ferdinand Broili (1874–1946).
Materials and methods
Slides with petrographic thin-sections
Nine slides with petrographic thin-sections of ichthyosaur
teeth were identified as being part of the doctoral thesis
materials of Bauer (1898). Previous to this date, the
Ichthyosaur tooth thin-sections
collection of Upper Jurassic ichthyosaurs from Bavarian
lithographic limestone quarries in the ‘Bayerische Palaon-
tologische Staatssammlung’ (now BSPG) was thought to
have been completely destroyed during WWII (Dehm
1978).
Eight of the nine slides (Figs. 1a1–a8, 2a) belong to a
single ichthyosaur tooth (BSPG AS XIX 504a–h), which
was sectioned both transversally through the crown (serial
sections: slides 1–7, Fig. 1a1–a7) and longitudinally
through the root base (slide originally without number, here
referred to as slide ‘‘8’’, Figs. 1b, 2b, 4). In the original
caption to plate 26, Bauer (1898) noted that the transverse
sections seen in figures 28–30 belong to a tooth from the
Oberndorfer specimen, which is part of the syntype mate-
rial of Ichthyosaurus leptospondylus WAGNER, 1853.
Following the discussion of Bardet and Fernandez 2000,
the longitudinal sections would now be referable to Aeg-
irosaurus leptospondylus.
The remaining slide (Figs. 1b, 2b) shows an ichthyosaur
tooth (BSPG AS I 1656) longitudinally sectioned from
crown to root. The plane of section lays slightly parasag-
ittally, thus the cusp of the crown is missing. Bauer (1898,
p. 285) mentioned that figure 27 of plate 26 shows a lon-
gitudinal section of a tooth belonging to the Solnhofen
specimen, which was excavated at the Maxbruch, Solnhofen,
in 1894. According to Bardet and Fernandez (2000), this tooth
would be referable to an indeterminate ichthyosaur close to
the ophthalmosaurid Nannopterygius.
To sum up, BSPG AS XIX 504a–h belongs to the
Oberndorfer specimen = Aegirosaurus leptospondylus;
whereas BSPG AS I 1656 belongs to the Solnhofen
specimen = Nannopterygius.
Analysis of the slides
The slides were first photographed with a Nikon D2x with
Nikkor 105 mm micro lens (Fig. 1). The tooth microstructure
was then analysed in normal transmitted and polarized light
using a Leica DM2500 M composite microscope mounted
with a Leica DFC 420 C digital camera.
Results and discussion
Matching slides and original drawings
Due to the level of detail (especially of the pattern of
micro-cracks in the histological tissues) in the drawings of
Bauer (1898), it was possible to relocate and match the
exact areas (Fig. 3) in the original images of figure 28–30
of plate 26: figure 28 shows part of slide 6; figure 29 part
of slide 5 and figure 30 part of slide 4. The planes of
Fig. 2 Overview of
microscopic images of all
sections. a1–a8 BSPG AS XIX
504a–h (slides 1–7 and ‘‘8’’),
tooth from the Oberndorfer
specimen, referable to
Aegirosaurus leptospondylus,
sectioned transversely and
longitudinally. Note resorption
pit and remnant of enamel
probably belonging to a
replacement tooth at lower right
margin of root in a8. b BSPG
AS I 1656, longitudinal section
of a tooth from the Solnhofen
specimen, referable to
Ichthyosauria indet. aff.
Nannopterygius
T. M. Scheyer, M. Moser
sectioning of the figured slides are thus situated at mid-
crown height (figures 29, 30) and at the neck of the tooth
(figure 28). As such, we can be positively sure that these
slides were indeed used to create the original figures. In all
three slides, the shown orthodentine and outer enamel layer
is in various stages of deterioration due to borings of
Mycelites ossifragus. Furthermore, it is plausible to assume
that the slides 2 and 3 are those Bauer (1898, p. 287)
referred to when mentioning the presence of ‘‘Furchen’’
(=grooves) in two cross sections, which were apparently
cut 2.0 and 3.0 mm below the tip of the crown of a small
ichthyosaur tooth.
The slide with the longitudinal section (Fig. 4) is more
difficult to match with the original figure 27 of plate 26 of
Bauer (1898), for the following reasons:
1. The image of the sectioned tooth in the figure does
not express the same level of detail as seen in
figures 28–30 due to the lower level of magnification.
2. The image presents a tooth sectioned exactly through
the tip of the crown complete with undamaged borders
of the tissues.
3. The image lacks any indication of micro-cracks,
changes in coloration of the orthodentine, the enamel
Fig. 3 Matching slides 4–6
(BSPG AS XIX 504d–f) with
figs. 28–30 of plate 26 of Bauer
(1898); (fig. 28 = slide 6, no.
504f—fig. 29 = slide 5, no.
504e—fig. 30 = slide 4, no.
504d). Note that the new
microscopic images (slides 4–6)
have been rotated 90� clockwise
for technical reasons
Ichthyosaur tooth thin-sections
or the cementum layers, as well as differences in
radicular and pulpal mineral infillings.
The slide on the other hand, shows a similar but not
congruent shape and distribution of tissues seen in the
image. Otherwise, the crown, as well as complete tissue
borders is not visible. The slide furthermore shows histo-
logical details such as a pattern of micro-cracks or changes
in coloration, which apparently have not been used in the
production of the original figure. There are two possibili-
ties: first, the slide is indeed the original on which figure 27
is based, in which case Bauer would have presented a
highly stylized, reconstructed depiction of this ichthyosaur
tooth, not uncommon at the time of lithographic printing;
second, the slide does indeed belong to the sectioned tooth
shown by Bauer, but is not the section on which figure 27
is based. In this case, we would have to assume the pres-
ence of another section on which the crown might have
been completely present, but which, unfortunately, was not
preserved. Because the present slide represents the lower
section of the tooth only, the second possibility, also less
appealing, cannot be ruled out completely.
The slides 1–7 and ‘‘8’’ (BSPG AS XIX 504a–h)
The tip of the crown (slide 1; Fig. 2a1) shows a concentric
circular core of orthodentine ringed with a thin layer of
enamel. Enamel striation is absent. The dentine tubules are
arranged perpendicular to the outer surface of the tooth; the
central tubules extend towards the tip of the crown,
whereas the tubules surrounding the central area show the
radial arrangement, as seen in the other transverse sections.
The junction between enamel and orthodentine is smooth
without any dentine infolding.
Slides 2–6 (Fig. 2a2–a6) show an oval shape of the
enamel and orthodentine. Coarse striation of the enamel is
present in slides 2 and 3. At the enamel-dentine junction a
hypomineralised interglobular zone of the orthodentine is
visible (Fig. 5a, b). Slides 3 and 5 also reveal the presence
of peculiar scattered vascular canals (vaguely reminiscent
of primary osteons in bone) in the peri-pulpal orthodentine
(Fig. 5c–f). The vascular canals, which appear to be
directly connected with the pulp cavity, are here interpreted
to have resulted from secondary deposition of pulpal os-
teodentine, leading to a decrease in size of the pulp cavity
in these slides. Based on the scarcity of material, the origin
of this osteodentine tissue remains unknown. The other
transverse slides did not show similar vascular canals.
Where preserved, slides 4–6 again show a smooth outer
enamel layer, as well as a smooth enamel-dentine junction.
In slide 7 (Fig. 2a7), a small central core of pulpal cellular
osteocementum is surrounded by the orthodentine, which
itself is surrounded by cellular osteocementum vascular-
ised by mostly radially oriented canals. At the dentine-
cementum junction (Fig. 5g, h) the hypomineralised layer,
i.e. the granular layer of Tomes (e.g. Tomes 1876; see also
Peyer 1968; Dean 2000), is present. Furthermore, a thin
layer of acellular cementum, recently identified for the first
time in ichthyosaur teeth, i.e., in Platypterygius australis
from the Cretaceous of Australia (Maxwell et al. 2011), is
also identifiable between the orthodentine and the outer
cellular osteocementum. Slide ‘‘8’’ (Fig. 2a8) shows the
remainder of the root of the tooth, i.e. dentine surrounded
by pulpal and external osteocementum and the thin layer of
acellular cementum, in longitudinal section. A resorption
pit and a small part of dentine, possibly of a replacement
tooth are visible at the lower right margin of the section.
The longitudinal section (BSPG AS I 1656)
On this slide (Figs. 1b, 2b) the root and the neck of the tooth
in longitudinal section is preserved. The orthodentine is
bordered by pulpal osteocementum and external osteoce-
mentum (Fig. 6a, b). As in the transverse sections, a thin
layer of acellular cementum lies between the orthodentine
and the external cellular cementum (Fig. 6b). The layer
extends from the proximal tips of orthodentine up to the
neck of the tooth, at which it is exposed as in P. australis
(Maxwell et al. 2011). The enamel and associated part of the
orthodentine of the crown are not preserved on the slide.
Fig. 4 Matching the longitudinal section (BSPG AS I 1656) (on theleft) with fig. 27 of plate 26 of Bauer (1898) (on the right)
T. M. Scheyer, M. Moser
The pulpal osteocementum reaches up to ca. 1/3 of the
pulp cavity. The vascularisation is higher than in the
external osteocementum layer. The fibrous arrangement of
the pulpal osteocementum (Fig. 6a) is more ordered (yel-
low and blue colours in polarised light with Lambda
compensator) towards the pulp cavity and grades into an
Fig. 5 Histological details of the transverse sections on slides 2, 3, 5,
7 (BSPG AS XIX 504b, c, e, g). a, c, f and g seen in normal
transmitted light, d, in polarised light and b, e and h in polarized light
with lambda compensator. a, b Close-up of striated enamel and
orthodentine (slide 2). Note interglobular zone of dentine at enamel–
dentine junction. c, d Central pulpal area of tooth showing vascular
canals in the orthodentine (slide 5). e Central area of tooth with
vascular canals surrounded by orthodentine (slide 3). f Close-up of
vascular canal seen in c (slide 5). Note how dentine tubules deviate
around the canal. g, h Close-up of the cementum–dentine junction at
the level of the neck of the tooth (slide 7). Note zones of acellular
cementum and the granular layer of Tomes. AC acellular cementum,
CC cellular cementum, DT dentine tubules, E enamel, GT granular
layer of Tomes, IGZ interglobular zone of orthodentine, ODorthodentine, VC vascular canals
Ichthyosaur tooth thin-sections
unordered diffuse tissue towards the root base (pink col-
our). Similarly the external osteocementum has an
unordered structure (pink colour). The external border of
both the pulpal and external osteocementum is irregular
due to cavities and canals. Furthermore, coarse parallel
Sharpey’s fibres are present in the external osteocementum
at the tooth neck (Fig. 6b), whereas they appear less
ordered and conspicuous in the root base. The Sharpey’s
fibres are the remnants of the non-mineralised periodontal
ligament anchoring the cementum to the periradicular bone
in the living animal.
Incremental growth in dentine
In both the transverse and longitudinal sections of both
teeth sectioned by Bauer (1898), incremental growth marks
of the orthodentine can be identified in polarized light
(Fig. 7), although they are often obscured by the dentine
tubules, micro-cracks or diagenetic colouration. In the
transverse sections, the markings appear as sub-circular
light and dark banded rings in polarised light, whereas they
appear as light and dark band lines in longitudinal sections,
paralleling the outer dentine border. The lines can be
separated into two sets (Fig. 7), a very fine short period line
set where the spaces between lines varies around 2–3 lm
(=0.002–0.003 mm) and a coarser, long period line set
where spaces vary around 15–20 lm (=0.015–0.02 mm).
This is in contrast to the findings of Maxwell et al. (2011),
Fig. 6 Histological details of the longitudinal section BSPG AS I
1656. Images in a and b are both seen in polarized light with lambda
compensator. a Complete view of the sectioned tooth. At the root the
orthodentine (blue colours) is embedded in vascularised pulpal and
external cellular cementum of the osteocementum type. Note the
differences in structural orientation between the ordered upper pulpal
cementum (yellow and blue colours) and the irregular external and
basal root cementum (pink colours). At the cementum–dentine
junction, a thin layer of acellular cementum is visible (yellow colour).
b Close-up of the orthodentine pillar. Coarse parallel Sharpey’s fibres
are visible in the external cellular osteocementum. The orthodentine
shows a weak banding often obscured by dark dentine tubules. The
pulp cavity is filled with sparitic calcite minerals. AC acellular
cementum, CC cellular cementum, DT dentine tubules, GT granular
layer of Tomes, OD orthodentine, P pulp cavity, SC sparitic calcite,
ShF Sharpey’s fibres
Fig. 7 Detail of the orthodentine showing incremental lines of
growth in polarised light in the longitudinal section (BSPG AS I 1656,
Ichthyosauria indet. aff. Nannopterygius). Two sets of incremental
lines are visible: the first set is composed of very fine dark and light
lines (indicated by arrow heads), interpreted as daily lines of von
Ebner. The second set consists of broader dark and light bands,
indicating longer time periods, i.e. Andresen lines. DB dark band,
DT dentine tubules, LB light band
T. M. Scheyer, M. Moser
who did not find evidence for incremental lines in the or-
thodentine of P. australis.
As discussed by Dean (1998, 2000) it is much debated
how to identify the incremental growth marks in dentine,
so that similarly spaced lines might be interpreted either as
long or short period lines by different authors. Dean (1998,
fig. 7) showed that in human dentine seven to eight short
period lines (i.e. daily lines von Ebner spaced\3 lm apart;
after von Ebner 1902, 1906) are situated between long
period lines of Andresen (after Andresen 1898), which are
spaced approximately 20 lm apart. In reptiles, Erickson
(1996a), when studying the dentine deposition in juvenile
Crocodylus mississippiensis using double fluorochrome
staining, found daily growth lines of von Ebner in the
crocodylian dentine similar to those found in most mam-
mals. Erickson (1996b: p. 14625) also described the
presence of lines of von Ebner in dinosaur teeth, and noted
that: ‘‘dinosaur incremental line widths revealed a range of
6–28 lm and mean values ranging from 10.1 to
19.8 lm…’’ with a ‘‘…general increase in incremental line
widths through ontogeny in the taxa for which multiple age
specimens were available’’, leading to calculated tooth
replacement rates in dinosaurs between 46 and 777 days.
Following Erickson’s (1996b) approach, Sereno et al.
(2007) and D’Emic et al. (2009) calculated even higher
tooth replacement rates for sauropod dinosaurs, based on
counts of lines of von Ebner. Both works, however, did not
explicitly discuss Dean’s (1998, 2000) articles and the
possibility that the growth lines found in the sauropod teeth
might also present long period lines instead of daily
increments, in which case the replacement rate in sauropod
dinosaurs would increase significantly. Based on the two
sets of lines found in Bauer’s (1898) slides, we follow
Dean (1998, 2000) in identifying the finer set of growth
lines as lines of von Ebner, whereas the coarser set of
growth lines (or rather bands) are tentatively referred to as
long period Andresen lines.
The shape of ichthyosaur teeth in cross-section
Bardet (1990), who analysed characteristic dental cross-
sections, indicated that a circular shape of the crown and a
quadrangular outline in the lower part of the tooth root
apparently unites all Cretaceous ichthyosaurs. Later,
Maxwell and Caldwell (2006) noted that not only the
Cretaceous Platypterygius and Maiaspondylus possess this
character, but also the Upper Jurassic Brachypterygius,
whereas the character state remained unknown in the
Upper Jurassic Aegirosaurus. Given the fact that the series
of transverse sections can be referred to Aegirosaurus
leptospondylus (sensu Bardet and Fernandez 2000), this
character can also be addressed in the serial transverse
sections on slides 1–7 and the longitudinal section of slide
‘‘8’’: the sections indicate that, while the crown has a cir-
cular outline, the root of the sectioned tooth did not have a
quadrangular but an oval outline.
Regarding the second tooth (possibly aff. Nannoptery-
gius, according to Bardet and Fernandez 2000), Bauer
(1898) discussed the sub-quadratic shape of the root (the
shape of the external cementum) and refers to his plate 26,
figure 27. Without knowing the original outer shape of the
second tooth, however, the preserved longitudinal section
does not yield the information necessary to address the
shape of the root for this taxon.
Conclusions
Despite their advanced age, the slides prepared for Bauer’s
(1898) dissertation, are in exceptionally good condition. All
histological features of the ichthyosaur teeth could be studied
in the slides and all slides were matched with the original
figures and descriptions of Bauer. Following Bardet and
Fernandez (2000), the transversely sectioned tooth (slides
1–‘‘8’’; BSPG AS XIX 504a–h) belongs to the ophthalmosaurid
Aegirosaurus leptospondylus, whereas the longitudinally sec-
tioned tooth is derived from an indeterminate ichthyosaur close
to the ophthalmosauridNannopterygius. Studying the slides with
up-to-date knowledge of ichthyosaur dentition and tooth his-
tology further led to the recognition of an exposed layer of
acellular cementum (sensu Maxwell et al. 2011) at the tooth
neck, and daily and long period growth lines in the orthodentine
in both specimens. Some slides of the tooth of A. leptospondylus
furthermore revealed a more oval than rectangular shape of the
root and the presence of peculiar vascular canals, interpreted as
secondary osteodentine deposition, in the peri-pulpal orthoden-
tine. It is therefore documented that dental characters of
advanced ichthyosaurs are taxonomically useful for distin-
guishing at least the genera.
Acknowledgments Winand Brinkmann, Heinz Furrer, Christian
Klug, Ingmar Werneburg and Jerome Gapany (all PIMUZ) are
thanked for their varied help in acquiring information and discussions.
Irene Schwendimann (Zivilstandsamt Zug, Switzerland), Franz Moser
(Kirchheim, Germany) and the Diozesanarchiv Eichstatt (Germany)
are thanked for help in further acquiring biographical data. We further
would like to express our gratitude to the two reviewers Nathalie
Bardet and Vivian de Buffrenil, as well as to the editors Michael
J. Benton and Daniel Marty. This work was partly funded by the
Swiss National Science Foundation (No. 31003A_127053 to TMS),
MM is supported by the Volkswagenstiftung.
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Ichthyosaur tooth thin-sections
1
Electronic Supplementary Material
Biography of Franz Bauer
Franz Xaver Bauer was born on April, 28th 1870 in Dollnstein, a village between Solnhofen
and Eichstätt, Middle Franconia, as son of the smallholder Jakob Bauer and Kreszenz Heim
(Diözesanarchiv Eichstätt 1866-1884). After visiting Elementary School and Middle School
he successfully completed the Catholic High School in Eichstätt in 1889. Until 1894 he
studied Theology there and graduated as a priest. The following year he worked as a local
pastor but then decided to begin studies in natural sciences at the Royal Ludwig-Maximilians
University of Munich with a focus on geology and palaeontology (Anonymous 1903c). His
supervisor became Prof. Karl Alfred von Zittel, the first conservator (director) of the Bavarian
State Collection for Palaeontology and Geology (BSPG). At that time, the palaeontological
collection and museum was housed in the "Wilhelminum", the old building of the Bavarian
Academy of Sciences, Neuhauser Straße 51, in the city centre of Munich (Dehm 1978). The
student Franz Bauer had a chamber in the neighbourhood in just five minutes walking
distance (Amtliches Verzeichnis des Personals der Lehrer, Beamten und Studierenden an der
königlich bayerischen Ludwig-Maximilians-Universität zu München, Sommersemester 1895:
44). In 1897 Franz Bauer finished his Ph. D. with magna cum laude based on a dissertation on
the then known ichthyosaur material from the Upper Jurassic Solnhofen Limestone (Bauer
1898; this material is the subject of the present contribution). Afterwards in danger of
unemployment, Franz Bauer had to take the job of a private teacher in Madrid for the family
of the former Spanish ambassador in Washington (Anonymous 1903c, Weber 1904).
However, in 1899 Ferdinand Broili, then assistant for mineralogy and geology at the
Chemical Department of the Royal Technical High School (later Technical University of
Munich TUM), left his position to become assistant at the BSPG (Peyer 1946; Dehm and
2
Schröder 1948) and the open position at TUM was offered to and filled with Franz Bauer.
Immediately he became productive with scientific contributions on anthropological (Bauer
1900a,d) and palaeontological topics (Bauer 1900c). Already in 1900 he finished his
habilitation thesis on a new ichthyosaur species from the Lower Jurassic of Bamberg (Bauer
1900b) and thereupon was appointed as associate professor at the Chemical Department. In
this position he offered courses, which were well visited, and especially numerous field
courses, in palaeontology and geology. He was also in charge of the geological collection,
which he constantly improved with instructive rock and fossil samples for educational
purposes (Bericht über die Königliche Technische Hochschule zu München für das
Studienjahr 1899-1900 ... 1902-1903; the collection unfortunately did not survive World War
II, Prof. H. Scholz oral comm. Feb.2011).
Somewhat surprising, during this period of his life as a natural scientist, Franz Bauer did not
abandon his interest and connections to theology and so some of his writings appeared in
theological journals. In an essay [transl.] "On the evolution of organic life with special
consideration of vertebrates", Bauer (1900f) – probably to the utmost embarrassment of some
of his fellow theologists – clearly took position in favour of the Darwinian theory (which he
obviously had accepted as undisputable truth not even necessary to discuss). In another essay,
Bauer (1900e) summarized the evidence for an older age of the Earth of probably several to
100 million years. In September 1900 he served as secretary of the section for mathematics
and natural sciences at the Fifth International Congress of Catholic Savants, which was held
in Munich under the presidency of the famous French geologist Albert de Lapparent.
In the years from 1900 to 1903, Bauer was mostly occupied with his duties as a lecturer, and
so publications during this time were limited to a number of abstracts and reviews (Bauer
1901a-e, 1902a-c), although he had several projects in progress (Weber 1904). He also
became a member of the Deutsche Geologische Gesellschaft (Zeitschrift der Deutschen
3
Geologischen Gesellschaft. 54, Mitgliederverzeichnis 1.1.1903).
In 1903, at the age of 33, Franz Bauer, now Second Order Assistant and Associate Professor
for Mineralogy and Geology of the Mineralogical Laboratory of the Chemical Department at
the Royal Technical High School of Munich, came to a sudden death by an unfortunate
accident (Anonymous 1903c,d): On 20th of June, he joined a group of nine of his students to
celebrate midsummer night on the top of the 1826 m high Risserkogel Mountain south of
Tegernsee (Bavaria). When the group arrived at the summit late in the night, Franz Bauer
happened to change his shoes to wear more comfortable slippers. Together with the group of
young people Franz Bauer sat around a bonfire before at 1 a.m. he stood up, made a few steps
aside and, probably due to the moisture of the grass, suddenly slipped over the steep north-
eastern mountain side and fell down a 150 meters. The students immediately undertook every
effort under risk of their own lives, but could only rescue the shattered corpse on the next day.
Franz Bauer's death was the subject of several short notes in newspapers (Anonymous 1903a-
d) and scientific journals (Anonymous 1903e,f). An official obituary (Weber 1904), although
with only few biographical details, shortly characterized Franz Bauer as a highly respected
colleague and proficient vigorous teacher with a bright potential. Biographical sketches were
published by Wolff (1905) and Quenstedt & Quenstedt (1938). The most proliferous
biographical source, however, proved to be two consecutive articles of the Münchner Zeitung
(Anonymous 1903c,d), which were obviously not known to other biographers.
Obituaries of Franz Bauer
Anonymous. (1903a). Todesfall. [Dr. Franz Bauer]. Allgemeine Zeitung, 1903 (Beilage 139):
536; (Verlag der Allgemeinen Zeitung) München. (23.6.1903).
Anonymous. (1903b). Unglück in den Bergen. [Dr. Franz Bauer]. Münchener Post, 17 (138):
6; München. (23.6.1903).
4
Anonymous. (1903c). Das Unglück auf dem Risserkogel. [Dr. Franz Bauer]. Münchener
Zeitung · Unabhängige Tageszeitung, 1903 (140): 4; München. (23.6.1903).
Anonymous. (1903d). Zu dem Absturz des Herrn Dr. Bauer. Münchener Zeitung ·
Unabhängige Tageszeitung, 1903(141): 3. München. (24.6.1903).
Anonymous. (1903e). Der Privatdozent der Geologie an der Techn. Hochschule in München
Dr. Franz Bauer verunglückte am 21. Juni d. J. auf einer Exkursion bei Tegernsee.
Centralblatt für Mineralogie, Geologie und Palaeontologie, 1903(12), 394. Stuttgart: E.
Schweizerbart. [7.1903].
Anonymous. (1903f). Dr. Franz Bauer [obituary]. Leopoldina, 39(8), 100. (Kaiserlich
Leopoldino-Carolinische Deutsche Akademie der Naturforscher) Halle a. S. (8.1903).
Quenstedt, W., & Quenstedt, A. (1938). Bauer, Franz. In K. Lambrecht, W. Quenstedt, & A.
Quenstedt (Eds.), Palaeontologi · Catalogus bio-bibliographicus; Fossilium Catalogus,
1: Animalia, Pars 72 (p. 25). s'Gravenhage: W. Junk. [495 pp.]
Weber, M. (1904). Franz Bauer. Bericht über die Königliche Technische Hochschule zu
München für das Studienjahr 1902-1903, Beilage Nekrologe, 6. München: F. Straub ·
Kgl. Techn. Hochschule.
Wolff, G. (1905). Bauer, Franz, Dr. phil., Privatdozent für Geologie und Paläontologie in der
Chem. Abteilung d.Techn. Hochschule in München [obituary]. Biographisches
Jahrbuch und Deutscher Nekrolog, 8 (1. Januar bis 31. Dezember 1903), col. 10*; (G.
Reimer) Berlin.
Other biographical sources
Dehm, R. & Schröder, J. (1948). Ferdinand Broili 1874-1946. Neues Jahrbuch für
Mineralogie, Geologie und Paläontologie, Monatshefte, 1945-1948, Abteilung B,
Geologie und Paläontologie, 1948(9-12), 257–271.
5
Diözesanarchiv Eichstätt (1866-1884). Taufmatrikel Dollnstein, Bd. 11 (Taufen 1866-1884),
168 (No. 29).
Peyer, B. (1946). Ferdinand Broili, 1874-1946. Verhandlungen der Schweizerischen
Naturforschenden Gesellschaft, 1946, 358–360.
Extended bibliography of Franz Bauer
Bauer, F. (1898). Die Ichthyosaurier des oberen weissen Jura. [Ph. D. thesis 11.3.1897].
Palaeontographica, 44(5-6), 283–328. [pls. 25-27; Stuttgart: E. Schweizerbart; 5.1898]
Bauer, F. (1900a). Über den Schwund der Diploe an einem Philippinenschädel. Anatomischer
Anzeiger [Centralblatt für die gesamte wissenschaftliche Anatomie], 17(2-3), 58–62. [1
fig.; Jena: G. Fischer; 15.1.1900]
Bauer, F. (1900b). Ichthyosaurus bambergensis nov. sp. Beschreibung einer neuen
Ichthyosaurus-Art aus dem oberen Lias vom Geisfeld nebst einigen vergleichend-
anatomischen Bemerkungen über den Schultergürtel. [Habilitiationsschrift]. Bericht der
naturforschenden Gesellschaft in Bamberg, 18, 1-56. [6 figs., 2 pls.; Bamberg]
Bauer, F. (1900c). Osteologische Notizen über Ichthyosaurier. Anatomischer Anzeiger ·
[Centralblatt für die gesamte wissenschaftliche Anatomie], 18(24), 574–588. [18 figs.;
Jena: G. Fischer; 31.12.1900]
Bauer, F. (1900d). Ueber Schädel von den Philippinen. Archiv für Anthropologie [Zeitschrift
für Naturgeschichte und Urgeschichte des Menschen], 27(1), 107–116. [3 figs.;
Braunschweig: Fr. Vieweg; Deutsche Gesellschaft für Anthropologie, Ethnologie und
Urgeschichte; 9.1900]
Bauer, F. (1900e). Geologische Zeiträume. Monatsblätter für den kathol. Religionsunterricht
an höheren Lehranstalten, 1(8), 229–236. [Köln: J. P. Bachem; 8.1900]
Bauer, F. (1900f). Die Entwickelung des organischen Lebens mit specieller Berücksichtigung
6
der Wirbeltiere. Monatsblätter für den kathol. Religionsunterricht an höheren
Lehranstalten, 1(9), 257–262. [Köln: J. P. Bachem; 9.1900]
Bauer, F. (1901a). [Review of] Spandel, E., Eine fossile Holothurie (Synaptareste aus den
oberoligocänen Cerithienschichten des Mainzer Beckens). Abh. der naturhist. Ges.
Nürnberg, Bd. XIII, 1900. Geologisches Zentralblatt, 1(21), 667. [Leipzig: Borntraeger;
1.11.1901]
Bauer, F. (1901b). [Abstract of] Bauer, Fr., Ichthyosaurus bambergensis nov. sp. -
Beschreibung einer neuen Ichthyosaurus-Art aus dem oberen Lias vom Geisfeld nebst
einigen vergleichend-anatomischen Bemerkungen über den Schultergürtel. XVIII. Ber.
d. naturforsch. Ges., Bamberg, 1900, 56 Seiten, 2 Tfln. u. 6 Fig. i. Text.
(Habilitationsschrift.). Geologisches Zentralblatt, 1(21), 670. [Leipzig: Borntraeger;
1.11.1901]
Bauer, F. (1901c). [Abstract of] Bauer, Fr., Osteologische Notizen über Ichthyosaurus. Mit 18
Abbildungen. Anatom. Anzeiger, XVIII. Bd., 1900. Geologisches Zentralblatt, 1(21),
670. [Leipzig: Borntraeger; 1.11.1901]
Bauer, F. (1901d). [Abstract of] Ichthyosaurus bambergensis nov. sp. [Berichte der
naturforschenden Gesellschaft zu Bamberg, 18]. – Akten des fünften Internationalen
Kongresses katholischer Gelehrten[r] zu München vom 24. bis 28. September 1900.
Compte rendu du Ve Congrès scientifique international des Catholiques, 1901, 438.
[München: Herder]
Bauer, F. (1901e). [Abstract of] Osteologische Notizen über Ichthyosaurier. [Anatomischer
Anzeiger, 18]. – Akten des fünften Internationalen Kongresses katholischer Gelehrten[r]
zu München vom 24. bis 28. September 1900. Compte rendu du Ve Congrès scientifique
international des Catholiques, 1901, 439-440. [München: Herder]
Bauer, F. (1902a). [Review of] Roger, Otto, Ueber Rhinoceros Goldfussi Kaup und die
7
anderen gleichzeitigen Rhinocerosarten. Mit 2 Tafeln. 34. Bericht d. Naturwiss. Vereins
f. Schwaben u. Neuburg, Augsburg, 1900, 70 S. Geologisches Zentralblatt, 2(2), 58–59.
[Leipzig: Borntraeger; 15.1.1902]
Bauer, F. (1902b). [Review of] Spandel, Erich, Die Foraminiferen des Permo-Karbon von
Hooser, Kansas, Nordamerika. Mit 10 Abbildungen. Festschr. z. Säkularfeier d. nat.
Ges. in Nürnberg, 1901. Geologisches Zentralblatt, 2(15), 476. [Leipzig: Borntraeger;
1.8.1902]
Bauer, F. (1902c). [Review of] Spandel, Erich, Untersuchungen an dem
Foraminiferengeschlecht Spiroplecta im Allgemeinen, und an Spiroplecta carinata
D'Orb. im Besonderen. Mit 6 Abbildungen. Festschr. z. Säkularfeier d. nat. Ges. in
Nürnberg 1901. Geologisches Zentralblatt, 2(15), 476. [Leipzig: Borntraeger; 1.8.1902]
Reviews of Bauer’s works
Huene, F. von (1903). [Review of] F. Bauer: Osteologische Notizen über Ichthyosaurier.
(Anatom. Anz. 18. 1900. 574–588. 18 Fig.). Neues Jahrbuch für Mineralogie, Geologie
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Keilhack, K. (1901). [Review of] Bauer, Fr., Osteologische Notizen über Ichthyosaurier.
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einer neuen Ichthyosaurus-Art aus dem oberen Lias von Gaisfeld nebst einigen
vergleichend anatomischen Bemerkungen über den Schultergürtel. (XVIII. Ber. d.
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Koken, E. (1901). [Review of] Fr. Bauer: Die Ichthyosaurier des oberen weissen Jura.
8
(Palaeontogr. 44. 1898. 283–328. 3 Taf.). Neues Jahrbuch für Mineralogie, Geologie
und Palaeontologie, 1901, II, Ref.(3), 476. [Stuttgart: E. Schweizerbart].