Cretaceous Research 112 (2020) 104425

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Cretaceous Research
journal homepage: www.elsevier.com/locate/CretRes

Cranial palaeopathologies in a Late Cretaceous mosasaur from the

Netherlands
Dylan Bastiaans a, b, c, d, *, Jeroen J.F. Kroll e, Dirk Cornelissen b, John W.M. Jagt b,
Anne S. Schulp b, c, d
a €t Zürich, Pala
Universita €ontologisches Institut und Museum, Karl-Schmid-Strasse 4, 8006 Zürich, Switzerland
b
Natuurhistorisch Museum Maastricht, De Bosquetplein 6-7, 6211 KJ Maastricht, the Netherlands
c
Naturalis Biodiversity Center, Darwingweg 2, 2333 CR Leiden, the Netherlands
d
Faculty of Geosciences, Utrecht University, Princetonlaan 8a, 3584 CB Utrecht, the Netherlands
e
MUMCþ, Department of Radiology, P. Debyelaan 25, 6229 HX Maastricht, the Netherlands

a r t i c l e i n f o a b s t r a c t

Article history: Here we describe multiple pathological skeletal elements in a specimen assigned to a globidensine
Received 25 September 2019 mosasaur as Prognathodon cf. sectorius. This individual, NHMM 2012 072, was recovered from the upper
Received in revised form Lixhe 3 Member (Gulpen Formation, upper Maastrichtian) near Maastricht, the Netherlands. In all
14 January 2020
likelihood, it was bitten in the snout by a large, possibly conspecific mosasaur e and survived this attack.
Accepted in revised form 12 February 2020
The specimen described here is among the very few with clear and unambiguous evidence of (very likely
Available online 28 February 2020
intraspecific) agonistic interactions amongst mosasaurs. Despite significant injuries, including partial
amputation of the premaxilla, this animal initially recuperated from the encounter, but the subsequent
Keywords:
Trauma
infectious processes as a result of this attack were still ongoing at the time of death. Radiological and
Palaeopathology morphological features suggest chronic osteomyelitis which led to loss of bone within the left maxilla,
Mosasaurs which probably hampered the ability to feed, potentially contributing to its demise. This case study il-
Osteomyelitis lustrates the potential of integrative three-dimensional approaches in palaeopathological studies to
Maastrichtian provide a much more comprehensive and detailed description of alterations and underlying physio-
Intraspecific agonistic behaviour logical processes.
© 2020 The Authors. Published by Elsevier Ltd. This is an open access article under the CC BY license
(http://creativecommons.org/licenses/by/4.0/).

1. Introduction 1990; Bell, 2010; Bell and Currie, 2010; Rothschild, 2013; Foth et al.,
2015; Anne  et al., 2015; Hedrick et al., 2016; Xing et al., 2018).
On rare occasions, diseases and injuries sustained during the However, research on pathologies in mosasaurs (extinct Late
lifetime of an organism can be preserved in the fossil record, in the Cretaceous marine squamates; Russell, 1967) took off early, as is
form of osseous lesions and exostoses (Moodie, 1917, 1923; demonstrated in papers by Mudge (1877), Dollo (1882) and Gaudry
Rothschild and Martin, 2006). Palaeopathology, or the study of (1890). Their (mostly descriptive) works were succeeded by
these records of disease and trauma in archaeology and palae- numerous reports on both traumatic (e.g. Meijer, 1985; Mulder,
ontology, may provide a better insight into disease acquisition, soft 1985; Bell and Martin, 1995; Martin and Bell, 1995; Lingham-
tissue anatomy, healing strategies and even behavioural aspects of Soliar, 1998, 2004; Everhart, 2008) and non-traumatic conditions
extant and extinct taxa (Moodie, 1917, 1923; Monastersky, 1989, (e.g. Rothschild and Martin, 1987, 1993, 2006; Martin and
1990; Rothschild and Martin, 2006; Reisz et al., 2011; Anne  et al., Rothschild, 1989; Monastersky, 1990; Rothschild and Tanke, 1992;
2015). Although many palaeopathological surveys have been un- Mulder, 2001; Schulp et al., 2004), prompting the first hypotheses
dertaken in dinosaurs and archosaurs in general, the majority of on their underlying behavioural implications.
squamates have remained largely understudied (e.g., Monastersky, Multiple pathological specimens of various mosasaur genera
have been recovered from the Maastrichtian type area (southeast
Netherlands, northeast Belgium); these provide a glimpse into the
€t Zürich, Pala€ontologisches Institut und
interactions and trophic relationships of these marine squamates
* Corresponding author. Universita
Museum, Karl-Schmid-Strasse 4, 8006 Zürich, Switzerland. that inhabited this latest Cretaceous ecosystem (e.g. Kruytzer, 1957;
E-mail address: dylan.bastiaans@pim.uzh.ch (D. Bastiaans). Mulder, 1985, 1999, 2001; Lingham-Soliar, 1998, 2004; Schulp et al.,

https://doi.org/10.1016/j.cretres.2020.104425
0195-6671/© 2020 The Authors. Published by Elsevier Ltd. This is an open access article under the CC BY license (http://creativecommons.org/licenses/by/4.0/).
2 D. Bastiaans et al. / Cretaceous Research 112 (2020) 104425

2004, 2006). This record has subsequently been augmented by new and classification of the various pathologies (e.g. traumatic, infec-
finds from other locations (e.g. Bell and Barnes, 2007; Everhart, tious, traumatic-infectious, developmental and idiopathic) and
2008, 2012; Einarsson et al., 2010). Many pathological studies rely their respective criteria, follow the guidelines as presented by
on external morphological comparisons only, despite the avail- Hanna (2002) and Foth et al. (2015).
ability of histological, isotopic, other geochemical and radiological
techniques (Straight et al., 2009; Peterson and Vittore, 2012; Anne  2.4. CT Scanning
et al., 2015; Hedrick et al., 2016). In the present contribution we
report on a palaeopathological case study of a recently discovered The dentaries were CT scanned using a Philips Brilliance Big Bore
specimen of mosasaur, assigned to Prognathodon cf. sectorius at the Universitair Medisch Centrum (UMC) Utrecht, the
(NHMM 2012 072) from the type area and illustrate opportunities Netherlands, at 1-mm slice thickness, 120 kV, tube current of
that are provided by modern day integrative approaches (e.g. CT 263 mA, and an exposure time of 4031 ms, rendering 363 axial
and 3D-visualizations). Inherently destructive palaeohistological slices in 512  512 pixels. Elements from the upper jaw were CT
approaches were not applied, because this particular specimen scanned using a Siemens Somatom Definition Flash scanner at the
represents a unique occurrence of a rare taxon. Maastricht University Medical Center (MUMCþ), Maastricht, the
Netherlands, at 600-mm slice thickness, 230 kV, tube current of
2. Materials and methods 600 mA, and an exposure time of 500 ms, rendering 465 axial slices
in 512  512 pixels. The resulting DICOM stacks were converted to
2.1. Institutional abbreviation 3D models using a combination of Avizo v. 8.1, Meshlab v.1.3.2 and
Polyworks workspace manager v12.1.23 at the Rheinische Frie-
NHMM: Natuurhistorisch Museum Maastricht, Maastricht, the drich-Wilhelms-Universita €t Bonn, Germany and Mimics v21.0 at
Netherlands. the Pala€ontologisches Institut und Museum Zürich, Switzerland.
The specimen of the Asian water monitor (Varanus salvator, PIMUZ
2.2. Material A/III 1493), originally from the Zoo Zürich, was euthanised after
contracting pododermatitis on the 14th of May 1981. It was sub-
The specimen described here, NHMM 2012 072 was discovered sequently macerated by Dr. Rieppel and taxidermist Schoch and
on September 17, 2012 by quarry operator Carlo Brauer. It originates assembled by Badertscher & W. Etter in 1981. The monitor
from the upper third of the flint-rich Lixhe 3 Member of the Gulpen currently part of the palaeontological teaching collection of the
Formation (c. 68,3 Ma; Keutgen, 2018) of the former ENCI Hei- Pala€ontologisches Institut und Museum (Zürich, Switserland) was
delbergCement Group quarry, located south of Maastricht (Fig. 1), CT scanned using a Nikon XT H 225 ST at the Irchel (central) campus
making it the oldest known identifiable skeletal mosasaur occur- of the University of Zürich (Switzerland), at 300-mm slice thickness,
rence from the Maastrichtian type locality known to date (Jagt and 195 kV, tube current of 36 mA, rendering 771 slices in 1314  2492
Jagt-Yazykova, 2012; Schulp and Jagt, 2013; Keutgen, 2018). Bone pixels with an isotropic voxel size of 77,643-mm. The resulting tiff
material from these lower strata is preserved in a fashion that stacks were converted to 3D models using Mimics Materialise
differs markedly from that of vertebrates found higher up in sec- Innovation Suite v. 21.0 at the Pala €ontologisches Institut und
tion. A major difference is the presence and subsequent oxidation of Museum (PIMUZ), Universita €t Zürich (Zürich, Switzerland).
pyrite and marcasite in the matrix and the fossil bone producing
sulphur oxides, which in turn react with the carbonates of the 3. Description
matrix, resulting in a thin gypsum veneer on the surface of the fossil
(Ritsema and Groenenberg, 1993). This gives the fossil a progres- In the following section the pathological features of NHMM
sively greyish appearance with time, and will require additional 2012 072 will be discussed on the basis of their unique morphology
curatorial attention in order to prevent chemical disintegration of in comparison to normal healthy bone.
the bones.
The disarticulated assemblage includes partial sections of the 3.1. External morphology - premaxilla
maxillae, dentaries, braincase and parietal (Fig. 2), as well as
various elements of the axial skeleton. Some bone surfaces are The anteriormost section of the premaxilla exhibits a rugose
heavily abraded, which is markedly different from what is generally surface texture that extends dorsomedially from the tooth bases.
encountered higher up in the section. The presence of shed teeth of Anterioventrally within this rugose-textured surface is a sub-
odontaspidid sharks, together with tooth marks on the right den- circular depression of roughly two centimetres in diameter (a in
tary surrounding the neural foramen between the fourth and fifth Fig. 3B). The rugose area is bordered dorsally by a small, yet distinct
tooth positions, suggest at least some degree of scavenging by ridge of smooth cortical bone extending laterally from the ante-
pointed-toothed organisms. riormost neurovascular canal of the premaxillary (b in Fig. 3A, B).
Although preparation is still ongoing, some inferences can be Projecting dorsally from the second and sole remaining left pre-
made based on the available skeletal material. Tooth morphology, maxillary tooth present, a small pea-like bony structure is visible (c
including a swollen tooth base, labiolingually flattened crowns and in Fig. 3A, B). An elongated lesion is seen on the left side of the
sharp carinae, suggest affinities with Prognathodon sectorius anteriormost premaxillary neurovascular foramen, interconnected
(Schulp and Jagt, 2013). Extrapolation of the alveolar spacing in a with the foramen ventrally and separated merely by a bridge of
jaw section yielded an estimated body length of around thirteen smooth bone dorsally (d in Fig. 3A).
metres. Together with the fused haemal arches, this enormous Situated predominantly posteriorly on the right side of the
length would suggest the ontogenetic age to be close to maximum premaxilla and right maxilla, a roughening of the bone surface can
attainable size (i.e. full maturity). be distinguished (e in Fig. 3A). Very little smooth cortical bone can
be differentiated in this area, in comparison to other abnormal
2.3. Terminology regions.
The premaxilla shows an elongate, healed lesion that is oriented
The terminology used here to describe the appearance and anteroposteriorly, starting near the left premaxillary-maxillary
underlying disease processes, as well as the manner of description suture, slightly to the left of the anteroposterior central line (f in
 D. Bastiaans et al. / Cretaceous Research 112 (2020) 104425 3

Fig. 1. Provenance area (A) and stratigraphic position (B) of Prognathodon cf. sectorius NHMM 2012 072.

Fig. 3A, B). Although posteriorly similar in appearance to the right numerous smaller pits, closely match those of the adjacent pre-
lateral premaxillary surface, i.e., with a rugose appearance, the maxilla in both extent and appearance.
anteriormost section of the elongated bony protrusion consists of a
smooth bone surface, only intermittently interrupted by tiny pits. 3.2. External morphology - maxilla
Along the lateral edge of the lesion a broadened section is bordered
by the left premaxilla-maxilla suture, displaying a heavily distorted Some evidence of reactive bone can also be distinguished near
surface texture with various depressions and of a distinctly pitted the base of the first maxillary tooth on the left side which is situated
nature. Laterally, three distinct oblong pits (g in Fig. 3A, B) on the ventrolaterally from the anteriormost oblong pit (h in Fig. 3B),
left maxilla, along with local depressions in the surface layer and enclosed by depressions anteriorly and posteriorly. The swollen
appearance of the tooth base differs from that of the less strongly
swollen premaxillary tooth base. This might, in part, be due to a
combination of taphonomic overprinting (i.e. deformation) and the
phylogenetically distinct appearance of the tooth base (i.e. ‘prog-
nathodont’ condition). The exostosial bone near the left premaxilla-
maxilla suture, together with evidence of necrosis and exostiosal
bone growth on the tip of the premaxilla, possibly related to
smaller gouges located more posteriorly, give the overall bone a
distinct appearance that diverges notably from the natural state.

3.3. External morphology - dentary

A callus (i in Fig. 4C, D) is present on the anteriormost section of

the left dentary. This callus is centrally located on the lateral side of
the dentary fragment, directly beneath the first tooth position. A
Fig. 2. Skull elements of Prognathodon cf. sectorius (NHMM 2012 072) preserved
shown in red for the left side and in grey for the right side. Skull outline adapted from
small depression is surrounded by a rugose area. In contrast to its
Dortangs et al. (2002). (For interpretation of the references to colour in this figure counterpart, the preserved part of the left dentary completely lacks
legend, the reader is referred to the Web version of this article.) neural foramina in the anterolateral side, and shows the
4 D. Bastiaans et al. / Cretaceous Research 112 (2020) 104425

directions (b in Fig. 3A, B). This radiolucent region is markedly

different from the region surrounding the two adjacent openings in
the anteriormost premaxillary (d in Fig. 3A, and its neurovascular
foramen), which shows bone of a normal radiological appearance.
This rugose area can be linked to the elongate lesion that ex-
tends from the elevated structure on its dorsal edge (d in Fig. 3A).
However, the bony bridge in between these two adjacent holes in
the anteriormost portion of the premaxilla is also visible, showing
normal density. The small protrusion located near the sole surviv-
ing premaxillary tooth on the left lateral side, near the suture, has a
density that is similar to the above-mentioned rugose structures (c
in Fig. 3A, B).
The large rugose area (e in Fig. 3A), primarily positioned right
laterally from the anteroposteriorly-oriented ridge on the pre-
maxillary, is recognised by external to internal loss of bone, with
little or no decrease in density from its natural state. This can
further be discerned from infectious processes, as no signs of bone
repair or reactive bone growth are observed. This homogeneous
erosional pattern of bone surface in the affected area is even pre-
sent on the antero-posteriorly-oriented reactive bony growth, as
well as other skeletal elements including the medial surface of the
lower jaw elements. As a result, this erosion-based rugosity might
be attributable to post-mortem bacterial erosion.
The radiodensity of the ridge-like protrusion (f in Fig. 3A, B) and
the left maxillary (g in Fig. 3A, B), is significantly lower in com-
parison to the surrounding regions, including the large aberrant
region on the right lateral premaxillary. The most pronounced
differences in density can be observed in the posterior two-thirds,
mainly concentrated around halfway through the elongated ante-
roposterior exostosis of the premaxillary, and the adjacent section
of the left maxillary. Furthermore, the bony architecture sur-
rounding the first maxillary tooth base seems to have been altered
as well, with emphasis on the small tubular radiolucencies
extending towards the tooth base and the ventral surface of the
maxillary.

3.5. Specific radiological appearance

The anterior premaxilla is more radiolucent when compared to

the section directly posterior to the anterior premaxillary neuro-
vascular foramen (Fig. 5B). In contrast, the external bone surface of
most of the premaxillary, is covered by a layer of thick, compact
bone of ten to twelve millimetres in thickness. The thickness of the
same layer in the anteriormost and posterior section (i.e. at the base
of the exostosial ridge) of the premaxillary is reduced to merely a
couple of millimetres at most. This does not appear to be an artifact
Fig. 3. Prognathodon cf. sectorius left maxilla and premaxilla of NHMM 2012 072 in (A)
of preservation, as little post-mortem deformation or compaction
dorsal and (B) left lateral view. The letters denote the various abnormalities visible on
the exterior of the elements of the upper jaw. Note the extensive damage to the
has occurred at this end, and only minor erosion occurred. In
anteriormost section of the premaxillary (aed) and to the area on the left lateral side transverse view, the ridge-like structure of the premaxillary shows
surrounding the premaxillary-maxillary suture (feh). The rugosity on the right lateral a similar pattern (Fig. 5C). The reduced thickness of the outer highly
side visible on the premaxillary (e) is most likely associated with post-mortem bacterial dense (cortical) bone layer is restricted to the area underneath the
erosion.
ridge and the left maxillary, suggesting a correlation between the
enlargement of the ventrally located foramina. The right dentary two. The cortical bone layer seems to have been unaffected on the
displays a small lesion anterolateral to the anteriormost tooth po- right side of the premaxillary. A small triangular-shaped concen-
sition and directly adjacent to a large neurovascular foramen (j in tration of less dense irregular bony material can be distinguished
Fig. 4A, B). underneath the ridge-like protrusion which can be traced along the
entire length of this exostosis.
Although difficult to recognize, at the first premaxillary tooth
3.4. Overall radiological appearance position, faint contours of the alveoli on both sides are present in
the radiographs. The alveolar cavity is largely unaffected at its
The rugose anteriormost section of the premaxillary is more natural margins, yet appears partially filled by new, highly vascu-
radiolucent compared to the areas of normal density directly poste- larised tissue (Fig. 5A).
riorly (Fig. 5). The former region is characterised by a rugose surface The most dramatic variations in radiolucency are visible near
area centred around a roughly 2-cm-wide depression (a in Fig. 3B) the oblong lesions of the maxillary and the depression lateral from
capped by a smooth dorsoventral growth extending laterally in both the suture line on the premaxillary.
 D. Bastiaans et al. / Cretaceous Research 112 (2020) 104425 5

Fig. 4. Prognathodon cf. sectorius NHMM 2012 072 anterior dentary sections. (A) Right dentary in right lateral (A) and ventral (B) view; left dentary in left lateral (C) and (ventral (D)
view, respectively. Notice the callus formation on the anteriormost section of the left dentary (i). When comparing the two dentary section, a striking difference is the complete lack
of neural foramina on the surface of the left dentary fragment (D). Furthermore, slight widening of the neurovascular canals, visible in ventral view, can be observed. However, this
could be due to preservation and preparation artifacts (D). A small, less conspicuous lesion can be identified in the anteriormost one-third of the right dentary fragment (j).

Particularly in the left maxilla, canals developed in the subsur- the two dorsal pairs of canals and the pair of horizontal neuro-
face of this irregular and distinctly less dense bony material (Fig. 6). vascular tubular structures, while the former display different
The extent, position and orientation of the patterns observed sug- shapes, sizes and orientation between one another. It is therefore
gest a common origin for all or most radiolucent areas, as these all highly unlikely that the maxillary canals are part of a structured
appear to be in comparable states of bone remodelling and display network of neurovascular tubules, as illustrated by the difference in
a comparable number of radiodensity changes. canal morphology as mentioned above, and the presence of addi-
Similar to the ridge-like exostosis on the premaxillary, which is tional canals extending from the lateral subsurface of the maxillary.
bordered by bone of normal density on its right lateral side, the The left dentary fragment has no obvious neural foramina on the
irregular surface concentrated around the oblong holes is also (antero)lateral surface and large ventral foramina (Fig. 4). The
bordered by normal (dense) bone on its left lateral side. The area of radiological appearance (especially in coronal view) of the left
reduced density that extends posteroventrally from the oblong dentary fragment, however, displays a normal neurovascular ar-
holes on the maxillary, connects to the maxillary tooth base dor- chitecture with the anterioventrally projecting canals connecting to
solingually. The latter is indicated by an area of reduced density at the foramina visible on the outer surface (Fig. 7). The regularly
this end of the tooth base in comparison to its anterior side. spaced neural branching pattern is similar to the condition in the
The distribution of the canals underlying the pitted maxillary is opposite dentary, with dorsal offshoots connecting to the tooth
best observed in coronal view (Fig. 6A). The lesions on the exterior margins and anterioventrally to the foraminal canals. The thickness
of the bone are interconnected, forming a single central canal that and spacing of these canals is nearly identical in both dentary
runs anteroposteriorly throughout the maxillary (Fig. 6B, C). The fragments, suggesting that the external damage visible concerning
internal bony architecture enclosing this large tubular maxillary the widening of the foramina is most likely non pathological in
network differs in appearance from the sharply defined structures nature (i.e. preparation or preservation artifacts).
visible in the neurovascular architecture of the premaxillary. The bony callus that extends from the lateral surface of the left
Furthermore, the latter displays a highly organised interaction of dentary fragment, shows a distinct radiolucent cleft in transverse
6 D. Bastiaans et al. / Cretaceous Research 112 (2020) 104425

Fig. 5. Prognathodon cf. sectorius NHMM 2012 072 anterior segments of both maxillae. Clockwise from left to right: a 3D reconstruction using accompanied viewer software; (A) CT-
slice in coronal view. Notice that the alveoli for the anteriormost premaxillary teeth are still present internally despite being absent externally (arrows); (B) CT-slice in sagittal view,
showing the thickness of the outer bone layer (arrows), regularly interrupted by neurovascular channels; (C) CT-slice in axial view, illustrating the density changes underneath the
ridge-like structure located anteroposteriorly along the premaxilla-maxilla suture (arrow). Notice that the bacterial erosion on the right lateral side of the premaxillary has not
resulted in (significant) density differences. Furthermore, the severe decrease in the density is restricted to the left maxilla and adjacent section of the premaxilla and its ante-
riormost portion (B,C). The decreased density surrounding the right maxillary tooth can be explained by the presence of original limestone matrix.

view (Fig. 7A). The radiolucent cleft is centred in the bony exostosis, premaxilla and the oblong holes of the left maxilla (Fig. 9), provides
and extends nearly seven millimetres into the subsurface. The insight into the internal remodelling. The visualisation of the
positioning of exostosial growth and its architecture suggests that anomalous structure underlying the subsurface directly under-
the radiolucent cleft may in fact be related to the underlying cause neath the anteroposterior ridge-like exostosis, when compared to
of the damage to the dentary fragment. A second and much larger the adjacent normal cortical and subcortical bone layer, clearly
fracture with a wide spacing shows no sign of healing or reactive shows the common origin of these abnormalities, as no abnormal
bone growth. The large amount of remodelling and reactive growth irregular bony architecture was visible underneath the remainder
on the other fragments of the feeding apparatus, makes it more of the premaxillary cortex.
likely to be a consequence of post-depositional fracturing. The The thickness of the cortical bone layer shows a distinct
semicircular radiolucency visible in transverse view, representing decrease from the right lateral side of the premaxillary to the left
the central neurovascular canal anteroposteriorly-oriented in the (Fig. 8A). The overall thickness of the cortical bone layer seems
dentaries, is surrounded by non-reactive and regularly structured consistent on the right side of the premaxillary up to reaching the
bony architecture. edge of the ridge-like structure, where it slightly thickens, before
decreasing significantly on the left lateral side of the same struc-
3.6. 3D-modelling of pathologies ture, enclosed by another thickened section before reducing again
significantly on the maxillary. This clearly shows that the focus of
Three-dimensional visualisation of the dense external (cortical) this pathological bone remodelling was centred around the left
bone layer (Fig. 8A, B), the infill of the triangular-shaped irregu- slope of the ridge-like growth, and the oblong holes in the maxil-
larity underneath the exostosial ridge on the premaxilla (Fig. 8C, D), lary. A virtual infill of the triangular anomaly that underlies the
and the infill of the canals underlying both the foramina of the premaxillary ridge visualises the extent and shape of the reactive
 D. Bastiaans et al. / Cretaceous Research 112 (2020) 104425 7

Fig. 6. cf. Prognathodon cf. sectorius NHMM 2012 072 anterior upper jaw fragments, in (A) coronal, (B) sagittal, and (C) axial view. Channels visible extending anteroposteriorly and
dorsoventrally coagulating into a larger canal (arrows A-B, and right arrow C). Notice the difference between the neurovascular canals of the figure above in which the borders are
sharply defined, and those of the larger catchment area of the maxillary canals in this figure, which are rather smudged. These maxillary canals display both different shapes, sizes
and orientation when compared to the regular neurovascular cavities. The left arrow in (C) indicates the triangular radiolucency underlying the ridge-like structure.

bone (Fig. 8C, D). This subcortical structure extends posteriorly extending offshoots connecting to the neural foramina visible on
throughout the premaxillary, only to be restricted by the edge of the external surface of the premaxilla. These (central) tubules are
the fossil as preserved. The maximum subcortical extent of the parallel to one another, the distance in between the dorsal exten-
irregular architecture located anteriorly is nearly 2.5 cm and shal- sions are more or less equal, the width of each canal appearing
lows posteriorly to just over 1 cm at its posterior edge. consistent throughout their anteroposterior extent, and the two
The lateral extent of this anomalous structure, as measured, is sets of tubules do not interconnect with their counterparts into one
roughly 18e22 mm; dorsoventrally, this is only near 9 mm. central canal as is the case in the subcortical maxillary canals. The
In ventral view (Fig. 8C), the contours of the anomaly show two set of maxillary tubules has a significantly different overall archi-
sharp ridges parallel to one another and separated only by a minor tecture and size in comparison to the neurovascular structures
dip. These ridges become posteriorly less distinct. In general, lateral present in the premaxillary. The three dorsolaterally extending
broadening is observed dorsally before reducing again, producing a tubules, connected to the three oblong holes present on the exte-
very rough diamond-shape (Fig. 8D). The overall anomaly is ori- rior, have diverging widths and lengths. Major structural differ-
ented laterally to the left, under an angle of 50e70 to the dorso- ences are: (1) size differences of the individual maxillary canals; (2)
ventral plane (Fig 8A). the three maxillary canals all converge into a central canal, that
By comparing the tubular structures in the subsurface beneath measures more than three times the size of the neurovascular
the oblong holes in the left maxillary (Fig. 9B) with the regularly- structures; (3) the dorsolateral tubules all coalesce laterally from
spaced neurovascular canals (Fig. 9C), the underlying nature of different angles and at different positions to the large central cavity,
the former can be resolved. The vascular network underlying the very much unlike the highly organised structure of the premaxil-
premaxillary is made up of two central tubules that run ante- lary neuroarchitecture. An additional zone of radiolucency sur-
roposteriorly, regularly interrupted by dorsally and anterodorsally rounding the first maxillary tooth base is identified, which was
8 D. Bastiaans et al. / Cretaceous Research 112 (2020) 104425

Fig. 7. cf. Prognathodon cf. sectorius CT slices of right (A) and left (B, C) dentary fragments NHMM 2012 072, in sagittal (A, C) and coronal (B) view. (B) Notice the radiolucent cleft
centered in the protruding bony callus on the lateral surface of the dentary extending roughly seven millimeters into the bone (arrow). The semicircular radiolucency internally
represents the neurovascular canals indicated in (C). The adjacent fracture lines most likely represent post-depositional damage (B). Remnants of the anteriormost tooth are visible
dorsally in the sagittal view of the left dentary fragment, together with the neurovascular channels with the foramina projecting anteroventrally (C). For comparison (A), the regular
structure of the branching pattern of the right dentary fragment, extending dorsally towards the teeth and anteroventrally to the foramina on the anteriormost exterior surface.

externally enclosed by irregular surface textures, and seems to be Since most of the damage is located on the left side of the skull,
connected to the central canal of the maxillary. These features and the majority of the post-mortem (e.g. bacterial) erosion is
suggest that the structures visible in the subsurface of the left restricted to the right lateral side, it is therefore unlikely to have
maxillary do not represent neurovascular structures. Although been the underlying cause of the density loss on the premaxillary
comparisons to extant savannah monitor lizards seem to corrobo- and left maxillary. Furthermore, the dense compact bone is still
rate the absence of neurovascular structures with external present even in areas on the right lateral side that were subject to
foramina in this area in the “normal” non-pathological condition, much (bacterial) erosion. This indicates that bacterial erosion could
the possibility of these representing modified pre-existing neuro- not have been the cause of the large quantity of dense bony ma-
vascular structures cannot be ruled out (Fig. 10). terial on the left lateral side of the skull.

4. Discussion 4.2. Trauma: bite mark?

4.1. Distribution of lesions: pathological vs non-pathological The shape of the anteroposteriorly oriented ridge on the pre-
maxilla is reminiscent of a tooth strike lesion (Fig. 8C), similar to
The similarities in extent, location and degree of healing of the what has been reported in extant crocodilians and late Mesozoic
anomalous bony structures on the anterior side of the upper jaw theropod dinosaurs (Tanke and Currie, 1998, . 4; Drumheller &
strongly suggest a common origin. The extensive damage to the Brochu, 2004, 2-3). The shape and size of the subsurface anomaly
anteriormost section of the premaxillary is characterised by sig- that underlies this ridge, resembles a point of entry by the deepest
nificant loss of bone density, in comparison to the other sur- surface penetration and a shallowing upwards at the other end. The
rounding structures. Although post-mortem erosion and differential infill that was produced of this anomaly displayed on its ventral
mineralisation in the fossilisation process cannot be entirely ruled surface a sharp bifurcating line, ragged edges, and a strong incli-
out as a possible cause, the elevated anterior edge of the premax- nation at its base. Together with the posterior shallowing of this
illary makes a traumatic nature likely. This ‘edge’ of the premaxil- anomaly, this suggests a sharp object having been forced into the
lary is possibly related to the 2-cm-wide depression centred in the compact bone from the left lateral side, under an angle of 50e70 to
rugose area anteriorly, and the lesion interconnected to the neural the dorsoventral plane, and to a depth of nearly 25 mm deep at its
canal on the dorsal surface. maximum penetration. Subsequently, this object would have been
 D. Bastiaans et al. / Cretaceous Research 112 (2020) 104425 9

Fig. 8. Prognathodon cf. sectorius cortical bone thickness of the premaxillary of NHMM 2012 072, (A) in axial view. Notice variation in the thickness of the outer highly dense bone
layer from the right lateral edge of the premaxillary to the left, and the adjacent anteroposteriorly extending ridge-like exostosis. The area of reduced thickness is restricted to the
area underneath the ridge, suggesting that significant density changes and the exostosis share a common origin (cutoff value bone of normal density >900 HU, and abnormal bone
<<900 HU). (B) anteroventral view of the compact bone layer of NHMM 2012 072, and the infill of the triangular-shaped anomaly located underneath the ridge-like exostosis of the
premaxilla in (C above) right lateral, (C lower) ventral, and (D upper) anterior and (D lower) posterior view respectively. Notice that the subcortical structures are restricted to the
area underneath the premaxillary anteroposterior ridge. Furthermore, the thickness of the triangular infill is consistent with a point of entry or deeper surface penetration anteriorly
(C upper, right side) and a shallowing upwards posteriorly (C upper, left). In ventral view, a single or double central midline can be distinguished, and with the extent of the infill
becoming wider dorsally. This pattern too is very reminiscent of a drag mark, possibly involving a sharp object penetrating the cortical bone layer e.g. Peterson et al. (2009). Notice
that the dimensions and shape of the subcortical infill matches that of mosasaur teeth. More specifically, the dimensions correspond with teeth from a mosasaur that rivalled
specimen NHMM 2012 072 in size or possibly slight exceeded it.

dragged in a posterior direction (as it becomes progressively shal- reparative tissues indicates that the organism survived the initial
lower), projecting only several millimetres up to one centimetre facial injury for at least several weeks (Rothschild and Martin,
downwards below the normal cortical bone layer (the added height 2006; Xing et al., 2018).
of the ridge not taken into account). The dimensions of this lesion
correspond well to the size of an isolated tooth of the same indi- 4.4. Traumatic-infectious/bacterial infection and the mosasaur
vidual that is found associated, being little over one centimetre immune system
wide at the top, nearly twice that at the tooth base, and nearly
45 mm in length. This may be a good reflection of the size of the The oblong lesions on the maxillary and lateral side of the
dentition of the perpetrator. exostosial ridge on the premaxillary, together with the disrupted
bone surface with extensive pitting, might hint at a bacterial
4.3. Trauma: fracture infection subsequent to initial rostral trauma sustained. Extant and
fossil crocodylians, often used as an analogue for mosasaur
The bony callus, or periostal bone apposition on the lateral side behaviour, frequently display deep facial wounds that penetrate
of the left dentary fragment could be related to the damage on the bone, and those wounds often become a point of entrance for a
left side of the upper jaw. However, the origin of this type of variety of bacterial agents (e.g. Mackness and Sutton, 2000; Avilla
damage is rather ambiguous. A swelling or ‘callus’ is deposited by et al., 2004; Wolff et al., 2009; Martin, 2013). The vascular struc-
the periost, and may be a reaction to infection, cancer, irritation, or tures in the subsurface of the maxilla connecting the lesions into a
the result of a normal fracture-healing process (compare Schulp single wide canal, are not unlike the radiolucent endosteal changes
et al., 2004 for discussion in mosasaurs). This callus is very likely associated with the abscess cavities of pyogenic osteomyelitis (i.e.
the result of localised trauma, as a clear puncture can be seen in the pus-generating bone infection) as observed in mammals (And and
centre. Normal fracture healing involves the increase of periosteal Castello, 1995; Ortner and Putschar, 2003; Divers and Mader, 2005;
and medullary blood supply, followed by infilling of the fracture by Rothschild and Martin, 2006; Silverman, 2006; Jadwiszczak and
highly vascular fibrous callus. The exterior of the callus seems to Rothschild, 2019). Another indication for osteomyelitis with an
have started uniting the fractured surfaces with bony tissue, sug- exogenous cause might be the sclerotic margins lateral to the ridge
gesting a relatively advanced state of healing (Rothschild and and in-between the plate-like cortical bone structures of the pre-
Martin, 1993, 2006). The presence of a callus with various maxillary and maxillary respectively (Rothschild and Martin, 2006).
10 D. Bastiaans et al. / Cretaceous Research 112 (2020) 104425

Fig. 9. Prognathodon cf. sectorius NHMM 2012 072 (A); a 3D reconstruction of the infill underlying the three oblong lesions in dorsolateral view (B); and an infill of the regularly
spaced neurovascular canals present in the premaxilla (C). Notice the difference in both size and architecture. First off, the width differs significantly between the two structures.
Furthermore, the neurovascular structures all join dorsally into two central canals. The structures on the maxillary all join laterally into a single structure. The highly irregular
structures adjacent to the three maxillary canals surround the first maxillary tooth base. Together these features suggest that the structures in the maxillary do not represent
neurovascular structures or possibly modified pre-existing canals. (cut-off values were established with bone of normal density represented by > 800 HU; radiolucent abnormal
bone regions of <800 HU giving additional resolution required for a continuous, uninterrupted 3D representation).

The failure of the soft tissue and the newly produced bone to isolate Huchzermeyer and Cooper, 2000). What is interesting to note is
the infection likely led to extensive spreading, which initiated even that the tubular architecture of the maxillary lesions and the
more osteoblastic activity. The increased intraosseous pressure of suppurative-like appearance of the subcortex in the present
infection-induced inflammations may have interfered with mosasaur individual seems to be more closely comparable to pyo-
vascular supply, causing tissue necrosis and exposing underlying genic osteomyelitis of mammals than the granulomatous response
trabecular bone (Rothschild and Martin, 1993, 2006; Ortner and of reptiles. The infecting bacteria may have used pre-existing
Putschar, 2003). Although relatively little is known about the neurovascular structures similar to those existing in the anterior
innate and adaptive immune system of reptiles, and even contra- part of the premaxilla to spread and eventually drain the infectious
dictory observations have been made, their response to infection material, possibly even becoming chronic or systemic osteomye-
differs in some ways from that of mammals (Huchzermeyer and litis. On the other hand, some of these structures in the subsurface
Cooper, 2000; Silverman, 2006; Stacy and Pessie, 2007; may have been created solely by the pyogenic bacteria and thus
Zimmerman et al., 2010, p. 661). Overall, the secondary perios- represent osteolytic canals for draining the abscesses associated
teal, cortical and endosteal response to infection is less prominent with the large infectious area (e.g. as documented in Reisz et al.,
in reptiles than in mammals (Silverman, 2006). In fact, reptiles do 2011). The architecture of the subsurface structures in the maxil-
not produce liquid exudates (i.e. pus), which is a typical mamma- lary favours the latter explanation, as these are highly irregular in
lian response (Stacy and Pessie, 2007; Zimmerman et al., 2010). size and shape and do not display the same distribution of canals,
Instead reptiles generally produce solid aggregates of degenerated nor do they interconnect in a similar fashion as seen in neuro-
heterophils, termed caseous granulomas or fibriscesses, soon (<7 vascular structures. Furthermore, no canals in such orientation and
days) after extracellular infection, eventually turning chronic on the size have been identified by external observation of other (Prog-
long term (Huchzermeyer and Cooper, 2000; Stacy and Pessie, nathodon-like) mosasaurs, nor of any related extant taxa (collec-
2007; Zimmerman et al., 2010; Anne  et al., 2015). In short, when tions of NHMM, Fig. 10 here and e.g., Kass, 1999; Schulp et al., 2008;
triggered, the innate immune system of reptiles produces non- Lindgren and Schulp, 2010; Konishi et al., 2011; Grigoriev, 2013).
specific white blood cells (heterophilic leukocytes), which contain Although a degree of soft tissue involvement seems very likely, it is
the infection by forming a solid mass of ‘dead’ heterophils within unclear if these osteolytic canals were associated with extensive
the tissue (Huchzermeyer and Cooper, 2000; Stacy and Pessie, soft tissue abscesses or not.
2007; Zimmerman et al., 2010). The difference between hetero-
phils of extant reptiles and the roughly analogous neutrophils of 4.5. Traumatic origin: violent interactions?
mammalian immune systems is translated into the production of a
solid granuloma in the former and the production of fluid pus in the The shape of the anteroposteriorly oriented ridge-like lesion,
latter group (Stacy and Pessie, 2007; Zimmerman et al., 2010; the structure of the subcortical anomaly, the location and extent of
 D. Bastiaans et al. / Cretaceous Research 112 (2020) 104425 11

Fig. 10. Comparison of the premax-maxillary complex of cf. Prognathodon sectorius NHMM 2012 072, to that of extant monitor lizards. (A-B) the anterior skull morphology and
architecture as can be observed in the extant savannah monitor lizard (Varanus exanthematicus) and the Asian water monitor (Varanus salvator, PIMUZ A/III 1493) in right and left
lateral view, respectively. (C) radiograph (above) and three-dimensional reconstructions (D) in right lateral and left view (E) the same region visualized in the before-mentioned
species of Asian water monitor lizard in dorsal view. Notice, no multituberal structures with irregular spacing or distribution similar to those in NHMM 2012 072 representing
neurovascular canals can be distinguished in the maxillary or anterolateral exterior of the upper jawbones of the savannah monitor lizard (Varanus exanthematicus) and the Asian
water monitor (Varanus salvator). Seemingly confirming this observation, based upon the radiological imagery no endosteal analogues to the heavily branched tubular structures are
present in this species of extant monitor lizards. These observations concur with the diagnosis of these endosteal architectural features as reflecting the prolonged effects of chronic
pyogenic osteomyelitis. Data for the extant Varanus exanthematicus is from Schachner et al. (2013). Abbreviations: j ¼ jugal; l ¼ lacrimal; mx ¼ maxillary; pf ¼ prefrontal; and
pmx ¼ premaxillary.

the damage on both the maxillary, premaxillary and possibly the of the suspected tooth-strike lesion, the only plausible attacker
left dentary, together with the subsequent healing processes, all with sufficient bite force to penetrate the cortical bone to such an
indicate a traumatic origin of these pathologies, and are most extent is another mosasaur (Lingham-Soliar, 1998). Estimates of the
parsimoniously explained by a not immediately lethal interaction dimension of the aggressor seem to approach those of the victim, or
with one (or more) large mosasaur(s). Cranial lesions have previ- possibly even slightly exceed them, meaning in excess of ten me-
ously been noted in mosasaurs including reports from the Maas- tres. Extant crocodiles display a snout-grappling behaviour during
trichtian type section (Lingham-Soliar, 1998, 2004; Mulder, 1999; the mating season, while protecting territory, or in intraspecific
Schulp et al., 2004, 2006, 2013). Testifying to the aggressive in- competition over potential mates (Lingham-Soliar, 1995, 1998,
teractions between mosasaurs, a multitude of injuries have been 2004; Tanke and Currie, 1998; Peterson et al., 2009). This behav-
suggested as evidence, such as puncture wounds to the posterior iour has been postulated for mosasaurs in multiple cases of facial
side of the skull, anterior side of the upper jaw and lateral side of lesions (Bell and Gordon in Monastersky, 1989; Rothschild and
the mandibles (e.g. Williston, 1898; Bell and Martin, 1995; Martin, 1993, 2006; Bell and Martin, 1995; Lingham-Soliar, 1998,
Lingham-Soliar, 1998, 2004; Mulder, 1999; Schulp et al., 2004, 2004; Mulder, 1999; Schulp et al., 2004, 2006; Everhart, 2008) and
2006; Everhart, 2008). The earliest postulation for inter- or intra- a semi-lateral grappling attack would certainly explain the tooth-
specific behaviour in mosasaurs can be traced back to Williston strike mark and adjacent damage to the maxillary.
(1898, p. 214), who observed, ‘... exostosial growth in their lower
jaws, the vertebrae, especially those of the tail, and in the paddles, 4.6. Survival time
especially the digits.’’ The distribution of these exostosial growths
has been described in more detail in recent years, and can be The estimated survival time for this particular individual may
attributed to a variety of causes including infectious spondylitis, have ranged from a few weeks to a couple of months at least. The
diffuse idiopathic skeletal hyperostosis (DISH, or non-pathological extensive bone remodelling and the state of healing could have
‘tendon ossification’), traumatically-induced osteomyelitis, and occurred within several weeks after the sustained injury, as fibro-
normal fracture healing (Schulp et al., 2004, 2006). Direct and cartilagenous callus formation generally takes place within two or
especially unambiguous evidence of interactions between mosa- three weeks in humans and resorption occurs over a period of
saurs, however, is extremely rare and thus the present specimen sixteen weeks under normal conditions (Rothschild and Martin,
offers an extraordinary opportunity to reconstruct in more detail 1993, 2006; Lingham-Soliar, 2004). As far as bone repair is con-
one such interaction. cerned, a major difference between birds and mammals on the one
Considering the size of the victim, which is reconstructed to hand, and lizards and amphibians on the other concerning bone
have been well in excess of ten metres body length, and the shape repair, is that the latter do not produce secondary cartilage.
12 D. Bastiaans et al. / Cretaceous Research 112 (2020) 104425

Therefore, the bony callus present is not comprised of rapidly behaviour, extant crocodilians are often proposed. During mating
produced cartilaginous precursor material. Rather, endosteal new season, while protecting territory, or directly during intraspecific
bone forms slowly and is present after three weeks, setting a competition, extant crocodilians engage in display behaviour
relatively high minimum survival time for this organism (‘head-slapping’) or in grappling behaviour (Peterson and Vittore,
(Rothschild and Martin, 1993; Lingham-Soliar, 2004). However, the 2012). These encounters often result in serious injuries, even par-
rate of repair could be significantly decreased and thus the survival tial or complete amputations of limbs and jaw sections have been
time markedly extended, by the influence of infections or disease, noted, but such encounters are rarely lethal (Wolff et al., 2009).
as this is a reflection of the overall health of the organism Crocodilians and lizards have strong immune systems that enabled
(Lingham-Soliar, 2004). them to deal with the resulting infections (Lingham-Soliar, 2004).
The negative effects of the infection associated with the trauma The immune system of snakes, although representing largely un-
could have contributed to the eventual death of this individual. It explored terrain, seems to work largely similar to that of mammals,
appears that the infection was still ongoing at the time of death, although there are some important differences (Stacy and Pessie,
and possibly even starting to extend to the first maxillary tooth 2007; Zimmerman et al., 2010). Reptiles generally exhibit a stron-
base. ger innate system response (Stacy and Pessie, 2007; Zimmerman
et al., 2010). The innate immune response of ectotherms also pro-
5. Conclusions duces a body temperature increase, albeit not physiologically but
behaviourally, this makes the immune response of ectotherms
Specimen NHMM 2012 072 of Prognathodon cf. sectorius suf- relatively dependent on the environmental conditions (Stacy and
fered various injuries that are visible on the anterior sections of the Pessie, 2007; Zimmerman et al., 2010). Another difference is that,
premaxillary, left maxillary and left dentary, likely from an in contrast to mammals, the reptilian inflammatory response does
encounter with one or more mosasaur(s). Other large predators not produce a liquid pus exudate but rather a ‘cheese-like’ (i.e.
including sharks, rare crocodilians (e.g. Thoracosaurus and others) caseous) mass of fibrous tissues (a fibriscess) (Peterson and Vittore,
and elasmosaurids can be excluded or are deemed very unlikely as 2012; Zimmerman et al., 2010; Stacy and Pessie, 2007). Mosasaurs
based on the morphology of the lesions, their rare occurrence in the may further differ in that they may have reached a degree of
area, or the lack of adequate bite force (Mulder, 1998; Mulder et al., giganto- or endothermy, and therefore extant ectothermic reptil-
1998, 2000, 2016). The force of the trauma was concentrated on the ians may not present the ideal immunological models for these
left lateral side of the skull, approaching from an angle of 50e70 to marine reptiles.
the dorsoventral plane. This allowed for the teeth to penetrate the Mosasaurs would have had strong immune systems similar to
premaxillary and maxillary, as well as the left dentary at the same those of extant reptilians, as the extent of the injuries did not result
instance. It appears that one but likely more teeth dragged through in immediate death of this specimen. Interestingly, mosasaurs
the compact bone layer covering the premaxillary, resulting in an appear to differ in some respects as the canals in the subsurface of
anteroposteriorly-oriented gouge, later forming a ridge-like exos- the maxilla seem to hint at a liquid pus rather than typical reptilian
tosis. This may be due to movement of the victim relative to the caseous growths. Eventually the demise of this mosasaur seems to
attacker(s), resulting in multiple parallel scrapes in the subsurface have preceded the complete healing of these injuries, as reparative
of the premaxillary as the attacker's teeth dragged across its sur- processes were still ongoing at the time of death. The exact cause of
face. The concentrated force of the bite resulting in a localised death cannot be determined with certainty; however, since the
fracture in the left dentary, eventually leading to the formation of a infection seems to have still been ongoing at the time of death, the
reparative bony callus. Additional damage to the anterior part of the chronic infection and potentially reduced feeding capability may
premaxilla may also be indicative of a bite resulting in the ampu- have contributed to the eventual death of this animal. This case
tation of its anteriormost section, and obscuring of the first tooth study illustrates the great potential of integrative three-
position on both sides by bone reparative processes. Only portions dimensional approaches in palaeopathological studies to provide
of the premaxillary and maxillaries are preserved, the absence of a much more comprehensive and detailed description of alterations
the internarial bar and large portions of the maxillary obscure more and underlying physiological processes. Ideally pathological studies
details of this violent interaction. It is clear that this mosasaur should aim to incorporate both histological and radiological tech-
survived the encounter and very likely persisted for a significant niques (including 3D-visualizations) to supplement traditional
time. However the resulting infectious processes may have external morphological descriptions as suggested by previous au-
hampered the healing processes, while at the same time spreading thors (e.g. Straight et al., 2009; Peterson and Vittore, 2012; Anne 
throughout the premaxillary and maxillary. As a result, a vast et al., 2015; Hedrick et al., 2016). This allows for a much higher
amount of bone is dissolved in the maxillary, creating multiple degree of certainty in identifying a causative factor, as well as
anteroposteriorly oriented tubules and even starting to affect the providing a much more detailed description of skeletal processes
enamel of the first maxillary tooth base. The underlying nature of on a tissue level.
such an encounter between two (or more) proportionally similar
mosasaurs is highly speculative. However, the identification of
more such encounters in the fossil record, suggests either intra- or
interspecific competition or predation. According to the size of the CRediT authorship contribution statement
victim and the suggested attacker(s), the latter would appear
unlikely. Dylan Bastiaans: Conceptualization, Data curation, Formal
The Maastrichtian type locality is home to five species of four analysis, Investigation, Methodology, Software, Validation, Writing
different genera of mosasaur, including Carinodens belgicus (~3 m in - original draft, Writing - review & editing. Jeroen J.F. Kroll: Data
length), Plioplatecarpus marshi (~5e6 m), and the larger Progna- curation, Methodology, Validation, Resources, Writing - review &
thodon sectorius (>8 m), Prognathodon saturator (~12 m) and editing, Visualization. Dirk Cornelissen: Validation, Investigation,
Mosasaurus hoffmanni (14e17 m), although not all taxa have been Writing - review & editing. John W.M. Jagt: Validation, Resources,
reported from the lower stratigraphic reaches where NHMM Writing - review & editing, Supervision. Anne S. Schulp: Concep-
2012 072 was discovered (Mulder, 1999 and references therein; Jagt tualization, Validation, Resources, Writing - review & editing,
et al., 2008; Schulp et al., 2008, 2013). As an analogy for mosasaur Visualization, Supervision.
 D. Bastiaans et al. / Cretaceous Research 112 (2020) 104425 13

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no respect. http://oceansofkansas.com/mosapath.html.
Foth, C., Evers, S.W., Pabst, B., Mateus, O., Flisch, A., Patthey, M., Rauhut, O.W., 2015.
We owe many thanks to Ernst Smid and the radiology and New insights into the lifestyle of Allosaurus (Dinosauria: Theropoda) based on
radiotherapy department of the University Medical Centre Utrecht another specimen with multiple pathologies. PeerJ 3, 1e33 e940.
Gaudry, A., 1890. Les enchaînements du monde animal dans les temps ge ologiques:
(UMC, Utrecht, the Netherlands) and the forensic radiology
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Maastricht, the Netherlands) for providing the opportunity to scan sauridae). Proceedings of the Zoological Institute RAS 317, 246e261.
the dentary and the premaxillary-maxillary section of the upper Hanna, R.R., 2002. Multiple injury and infection in a sub-adult theropod dinosaur
Allosaurus fragilis with comparisons to allosaur pathology in the Cleveland-
jaw of the mosasaur, respectively, and helpful comments con- Lloyd dinosaur quarry collection. Journal of Vertebrate Paleontology 22,
cerning the diagnosis. Special thanks to Thorsten Plogschties 76e90.
(Rheinische Friedrich-Wilhelms-Universita €t Bonn) for help with Hedrick, B.P., Gao, C., Tumarkin-Deratzian, A.R., Shen, C., Holloway, J.L., Zhang, F.,
Hankenson, K.D., Liu, S., Anne , J., Dodson, P., 2016. An injured Psittacosaurus
the various three-dimensional reconstructions (Avizo, Meshlab and (Dinosauria: Ceratopsia) from the Yixian Formation (Liaoning, China): Impli-
Polyworks) and helpful feedback. Additional thanks to Bruce M. cations for Psittacosaurus Biology. The Anatomical Record 299, 897e906.
Rothschild, for new insights into and comments on the preliminary Huchzermeyer, F.W., Cooper, J.E., 2000. Fibriscess, not abscess, resulting from a
localised inflammatory response to infection in reptiles and birds. The Veteri-
work as presented during the 74th Annual Meeting in Berlin nary Record 147, 515e516.
(November 2014). DB was partially funded through the Swiss Na- Jadwiszczak, P., Rothschild, B.M., 2019. The first evidence of an infectious disease in
tional Science Foundation (grant no. 31003 A_179401 to T. Scheyer). early penguins. Historical Biology 31, 177e180.
Jagt, J.W.M., Jagt-Yazykova, E.A., 2012. Stratigraphy of the type Maastrichtian-a
We owe much gratitude to PD Dr. Torsten Scheyer
synthesis. Scripta Geologica - Special Issues 8, 5e32.
€ontologisches Institut und Museum, Universit€
(Pala at Zürich) and Jagt, J.W.M., Cornelissen, D., Mulder, E.W.A., Schulp, A.S., Severijns, J., Verding, L.,
Dr. Ashley Latimer for providing a scan of the Asian water monitor 2008. The youngest in situ record to date of Mosasaurus hoffmanni (Squamata,
(Varanus salvator). Furthermore, we are grateful to PD. Dr. Torsten Mosasauridae) from the Maastrichtian type area, the Netherlands. In:
Everhart, M. (Ed.), Proceedings of the Second Mosasaur Meeting, vol. 3. Fort
Scheyer and Louis Verding (NHMM, Maastricht, the Netherlands) Hays Studies Special Issue, pp. 73e80.
for helpful discussions on normal and pathological appearance in Kass, M.S., 1999. Prognathodon stadtmani:(Mosasauridae) a new species from the
histological and radiological sections. We would like to thank the Mancos Shale (lower Campanian) of western Colorado. Vertebrate Palae-
ontology in Utah 99e1, 275e294.
former ENCI HeidelbergCement Group for support with the exca- Keutgen, N., 2018. A bioclast-based astronomical timescale for the Maastrichtian in
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keen eye to spot and recognise the fossil and for persuading his Konishi, T., Brinkman, D., Massare, J.A., Caldwell, M.W., 2011. New exceptional
superiors to postpone quarrying operations at the site. Further- specimens of Prognathodon overtoni (Squamata, Mosasauridae) from the upper
more, without the dedication, professionalism and effort of all Campanian of Alberta, Canada, and the systematics and ecology of the genus.
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volunteers and others involved in preparation at the Natuurhis- Kruytzer, E.M., 1957. De Mosasaurus van Bemelen. Mosasaurus hoffmanni Mantell.
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Cretaceous, Upper Maastrichtian of The Netherlands. Philosophical Transactions
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