Received: 10 June 2024 Revised: 11 July 2024 Accepted: 27 July 2024

DOI: 10.1002/ar.25563

RESEARCH ARTICLE

Prey size and ecological separation in spinosaurid

theropods based on heterodonty and rostrum shape

Domenic C. D'Amore 1 | Evan Johnson-Ransom 2 | Eric Snively 3 |

David W. E. Hone 4

1
Department of Natural Sciences, Daemen
University, Amherst, New York, USA Abstract
2
Department of Organismal Biology and Members of the dinosaur clade Spinosauridae had numerous traits attributed
Anatomy, University of Chicago, Chicago, to feeding in or around water, and their feeding apparatus has often been con-
Illinois, USA
3
sidered analogous to modern crocodylians. Here we quantify the craniodental
Oklahoma State University College of
Osteopathic Medicine–Cherokee Nation,
morphology of Spinosauridae and compare it to modern Crocodylia. We mea-
Tahlequah, Oklahoma, USA sured from spinosaurid and crocodylian skeletal material the area of alveoli as
4
School of Biological and Behavioural a proxy for tooth size to determine size-heterodonty. Geometric morphometrics
Sciences, Queen Mary University of
were also conducted on tooth crowns and tooth bearing regions of the skull. Spi-
London, London, UK
nosaurids overall had relatively large alveoli, and both they, and crocodylians,
Correspondence had isolated regions of enlarged alveoli. Spinosaurines also had enlarged alveoli
Domenic C. D'Amore, Department of
along the caudal dentary that baryonychines lacked, which instead had numer-
Natural Sciences, Daemen University,
4382 Main Street, Amherst, NY, 14226, ous additional caudal tooth positions. Size-heterodonty was positively allometric,
USA. and spinosaurids overlapped with generalist/macro-generalist crocodylians of
Email: ddamore@[Link]
similar sizes. Spinosaurid crown shape morphologies overlapped with certain
slender-longirostrine crocodylians, yet lacked molariform distal crowns typical
of most crocodylians. Spinosaurid rostra and mandibles were relatively deep
with undulating margins correlating with local tooth sizes, which may indicate
a developmental constraint. Spinosaurines had a particularly long concavity cau-
dal to their rosette of anterior cranial teeth, with a corresponding bulbous rostral
dentary. The spinosaurid feeding apparatus was well suited for quickly striking
and creating deep punctures, but not cutting flesh or durophagy. The jaws inter-
locked to secure prey and move it deeper into the mouth. The baryonychines
probably did little oral processing, yet spinosaurines could have processed rela-
tively large vertebrates. Overall, there is no indication that spinosaurids were
restricted to fish or small aquatic prey.

KEYWORDS
Baryonychinae, crocodylia, dentition, feeding, heterodonty, jaw, rostrum, Spinosauridae,
Spinosaurinae, teeth

1 | INTRODUCTION mostly fragmentary remains with nevertheless distinct

diagnostic features such as extended dorsal neural spines,
Spinosaurids were an enigmatic and unusual group of large humeri and manual claws, and elongate, narrow
large theropods (Hone & Holtz, 2017). They consist of jaws with a terminal rosette (a rounded, tooth-bearing

Anat Rec. 2024;1–18. [Link]/journal/ar © 2024 American Association for Anatomy. 1

2 D'AMORE ET AL.

anterior expansion of the rostrum) (Hone & Holtz, 2017; consuming birds (Dessborn et al., 2011; Milardi
Holtz et al., 2004; Sereno et al., 1998; Sues et al., 2002) et al., 2022), brown trout eating rodents (Milardi
Their dentition was variable but crowns were generally et al., 2016), and even dramatic examples such as orcas
elongate and conical, with a reduction in the degree of eating moose (Jefferson et al., 1991).
labiolingual compression and tooth serrations that The first spinosaurid discovered, Spinosaurus aegypti-
were common to other theropods (Hone et al., 2010; cus (Stromer, 1915), had a partial mandible but no cra-
Kellner & Campos, 1996). Spinosaurids may be split nium or rostrum, and was reconstructed as having a
into two major clades. These are similar overall but skull similar to other large theropods (Stromer, 1936).
show clear differences in anatomy, including in their With the discovery of Baryonyx walkerii (Charig &
jaws (Hone & Holtz, 2017; Lacerda et al., 2022; Sales & Milner, 1986) and Suchomimus tenerensis (Sereno et al.,
Schultz, 2017). Spinosaurinae generally consisted of 1998), it became increasingly apparent that the spino-
larger members and had a dorsal sail and fewer tooth saurid skull represented a derived condition that deviated
positions, with the Baronychinae having been smaller, from the deeper, shorter rostrum seen in more ‘typical’
lacking said sail (though they did have dorsal and theropods (demonstrated in Rayfield, 2011). Comparisons
sacral neural spines longer than most other theropods), have been made to phytosaurs and pike eels (Vullo
and had more numerous teeth (Sereno et al., 1998). et al., 2016; Yun, 2024), but the most common analogy is
Although numerous isolated spinosaurid teeth have with modern Crocodylia (Holtz, 1998), as exemplified by
been recovered, very few have been found in situ. Suchomimus which literally translates to ‘crocodile
Extensive arguments persist about spinosaurid life- mimic’ (Sereno et al., 1998). The crocodylian feeding appa-
styles, especially the degree to which they were aquatic. ratus shares several traits with spinosaurids, such as a long
Bone histology indicates aquatic affinities to a certain upper-jaw with a secondary palate, an undulating or fes-
degree (Aureliano et al., 2018). Although evidence of a tooning jaw margin, and often a rosette (Brochu, 1997,
diverse diet exists, including terrestrial prey based on iso- 1999, 2001; Cuff & Rayfield, 2013; Rayfield et al., 2007). The
topic data (Amiot et al., 2010), gut contents (Charig & shape of this snout is often correlated with diet. Drumheller
Milner, 1997), and embedded teeth (Buffetaut et al., 2004), and Wilberg (2020), for example, categorized all modern
it is still likely fish were a major part of their diet. Addi- crocodylians into three ecomorphotypic groups based on
tionally, most raptorial marine amniote teeth are conical rostrum shape and prey size. These are: (1) “slender longir-
(Fischer et al., 2022) like those of spinosaurids. Ibrahim ostrine” with slender- and tube-snouted taxa that take prey
et al. (2014, 2020) spearheaded the idea Spinosaurus was a smaller than themselves, (2) “generalists” with mesoros-
subaqueous pursuit predator with tail propelled locomo- trine, triangular snouts and consuming prey around their
tion and potentially reduced terrestrial mobility (see also own body mass, and (3) “macro-generalists” with mesoros-
Amiot et al., 2010; Arden et al., 2019; Fabbri et al., 2022). trine, U-shaped snouts that eat prey larger than themselves.
This scenario has been met with resistance, as others have Crocodylian teeth are also similar to those of spinosaurids
called into question the animal's stability in water, hydro- (Hone & Holtz, 2017), and cluster together concerning lin-
dynamics, and the degree to which its anatomy was spe- ear morphometrics (Navarro-Lorbés & Torices, 2018). Cro-
cialized for an aquatic lifestyle (Henderson, 2018; Hone & codylians typically have oscillating size heterodonty of the
Holtz, 2019; Myhrvold et al., 2024; Sereno et al., 2022). teeth in the jaw, but linear shape heterodonty, with
Alternative hypotheses include preponderance as a shore- crowns changing from relatively caniniform (acute and
line or wading feeder that hunt from above the water's sur- pointed) to molariform (stout and bulbous) with more
face (Hone & Holtz, 2021), which has been further distal positions (D'Amore et al., 2019). Similar to snout
supported by skull morphology and nares position shape, tooth morphology is linked to prey size, as well
(Smart & Sakamoto, 2024). However, despite the contro- as degree of processing and prey hardness
versy, to date there has been little discussion on potential (Brochu, 2001; D'Amore et al., 2019).
prey types and how they might have been acquired. This is Despite that the spinosaurid feeding apparatus has
especially true relative to certain other theropod groups been described numerous times and the similarities to
where more direct evidence of trophic interactions is avail- crocodylians often noted, few studies have conducted rig-
able (DePalma et al., 2013; Erickson & Olson, 1996; orous, quantitative comparisons between the two. Ray-
Longrich et al., 2010; Peterson & Daus, 2019; Rogers field et al. (2007) and Cuff and Rayfield (2013) created 3D
et al., 2007; Sakamoto, 2022; Therrien et al., 2023). Even models and compared the structural properties of select
animals that are very aquatic can take primarily terrestrial spinosaurids of both subgroups to crocodylians, but there
animals depending on the context and opportunities. have been no other quantitative comparisons. Dentition
Examples include terrestrial mammals eaten by large cro- in spinosaurids has been described using both discrete
codylians (Wallace & Leslie, 2008), pike and catfish characters and linear morphometrics (e.g., Hendrickx
D'AMORE ET AL. 3

et al., 2015, 2019) but has usually only been compared all available alveoli from 16 crocodylian specimens (cov-
with other theropods. The purpose of this study is to con- ering 10 species) and 18 spinosaurid specimens (covering
duct an in-depth, quantitative analysis of craniodental 10 species with several designed as ‘indeterminate’) total-
morphology throughout Spinosauridae, and directly com- ing in 1346 alveoli. Images of specimens were uploaded
pare it to modern Crocodylia using similar data collection into TpsDig2.16 (Rohlf, 2021), and four landmarks were
techniques. We use a three-pronged approach, investigat- plotted along each alveolar margin at the mesial, distal,
ing tooth alveolar size, crown shape, and rostral dimen- labial, and lingual extremes. The straight line distances in
sions. Through analogy with modern crocodylian life millimeters between the mesial-distal and labial-lingual land-
histories, we elaborate upon spinosaurid fundamental marks were used as lengths and widths respectively, and an
niche characteristics. We also expand upon variability elliptical area was derived using the following equation:
within the group, explore the effects of allometry, and
determine what similarities and differences existed Alveolar area ¼ π  ðlength=2Þ  ðwidth=2Þ:
between Baryonychinae and Spinosaurinae.
Spinosaurid and crocodylian tooth crowns were typically
mesiodistally symmetrical at the base and filled much of
2 | MATERIALS AND METHODS their alveoli. Therefore equations for cross-sectional areas
that vary consistently from that of an ellipse
Spinosaurid fossil material was measured directly by the (Motani, 2001; Snively et al., 2013) would not affect rela-
authors from museum collections where possible, as well tive coronal areas of the teeth. Alveolar area was col-
as from the figures of published manuscripts where nec- lected from both the left and right sides when possible,
essary. Modern crocodylian data were all collected from and the values were averaged for the symmetrical posi-
museum specimens either photographed directly by the tions. A methodological exception was the single speci-
authors or taken from data published in D'Amore et al. men of Angaturama limai (USP GP 2T-5), which was
(2019). Institutional abbreviations include MSNM V4047 fractured through the alveoli along the mesial-distal
(Museo di Storia Naturale di Milano, Milan, Italy); NHM plane (Kellner & Campos, 1996). We instead measured
(Natural History Museum, London, United Kingdom); MNN the length of the alveolus along this plane and applied
(Musée National du Niger, Niamey, Republic of Niger); and this value for both length and width in the equation
USP (Universidade de São Paulo, São Paulo, Brazil). above to derive alveolar area.
Photographs were taken with multiple cameras with no Designating a size covariate that could be collected
wide angle, zoom, or macro settings (including a Canon from all spinosaurid individuals was not possible
EOS T3, Canon EOS 70D, and an iPhone 11) and a scale because they vary in completeness, so we used several.
was included. Concerning data taken from published fig- First, we calculated head width (HW), defined here as
ures, photographs were prioritized over line drawings the linear distance between the caudal-most junction at
and/or digital reconstructions whenever possible. Care was the quadrate and quadratojugals on the lateral margins
taken to ensure that published photographs were all taken of the skull. It was measured for all crocodylian speci-
at the same orientation for the purpose of consistency. Only mens, but could only be collected from reconstructions
‘large’ crocodylian taxa were considered, with species that of Irritator challenger and Suchomimus tenerensis (taken
rarely reach 3 m in length excluded. Therefore, any species from fig. 4 of Schade et al., 2023 and fig. 2 of Sereno
within Caiman, Osteolaemus, and Paleosuchus, as well as et al., 1998, respectively). Second, we calculated the
Alligator sinensis, were omitted. All remaining alligatorids maximum width at the premaxilla. The rostral-most
fell within the macro-generalist category, and all gavialids region of the upper jaw in spinosaurids forms a ‘termi-
fell with the slender-longirostrine category [Tomistoma nal rosette,’ and is a commonly preserved jaw feature
schlegii is considered a member of Gavialidae here (Pan (detailed in Lacerda et al., 2022). Referred to as rosette
et al., 2021)]. All crocodylids fell within the generalist cate- width (RW), this was also calculated for all crocody-
gory, except for Crocodylus johnstoni and Mecistops cata- lians except alligatorids as they do not have the rosette
phractus which were slender-longirostrine. See Tables S1 morpholotype.
and S2 for sources of all specimens. We measured both isolated and in situ crowns. Iso-
As archosaurs are thecodont with socketed teeth lated crowns are common in collections and made up the
(Edmund, 1969), we used alveolar dimensions as a proxy bulk of the sample for Spinosauridae, but a small amount
for tooth size. The dimension of interest was the area of of in situ crowns were also available. All crowns sampled
the alveolar opening, and was derived from images ori- from Crocodylia were in situ, most of which were taken
ented from the palatal and dorsal perspective of the cra- from D'Amore et al. (2019). In addition, tooth crowns of
nia and mandibles respectively. Data were collected for the megalosaurid sister clade to Spinosauridae were
4 D'AMORE ET AL.

added to the dataset to compare with the more derived A measure of heterodonty was calculated using the
dental condition seen in Spinosauridae. This resulted in a formula for morphological disparity derived by Foote
total of 732 crowns. Images used for data collection were (1993) (see also Sheets & Zelditch, 2013; Zelditch
from the labial perspective of the tooth perpendicular to et al., 2003) used in past heterodonty studies
its long axis. Teeth with worn apices were included, but  P m 2015;
(D'Amore,  D'Amore et al., 2019) defined as
we omitted any teeth with large wear facets or chips. We MD ¼ i¼1 i =ðm  1Þ. Morphological disparity (MD)
D 2

also omitted teeth if a crack occurred that distorted the is the sum of the differences of the values of a given alve-
shape of the tooth. olar area (i) from the mean for the entire specimen (Di)
We outlined each tooth crown by tracing its edges. squared, with the number of alveolar positions (m)
Images were entered in TpsDig, and the tooth was traced factored in.
using the curve drawing tool along the enamel margin. Alveolar areas and heterodonty were used as depen-
This resulted in two curves along the mesial and distal dent variables in a series of regression analyses. Average
sides. TpsDig then transformed each curve into alveolar area (AAA) was the average of area of each alve-
30 equidistant coordinates, and we then combined the olus for all tooth positions sampled. We regressed AAA
apical-most coordinates. This resulted in three discrete for each specimen against HW and RW using reduced
landmarks (two at the base and one at the apex) and major axes regressions in PAST (Hammer, 2001). We next
56 semilandmarks. This coordinate number has been plotted disparity (MD) against AAA, HW, and RW also
used in numerous prior studies (D'Amore, 2015; D'Amore using reduced major axes. The goals of these were to (1)
et al., 2018, 2019; Smith et al., 2005; Tinius & Patrick determine if the dependent variables increased allometri-
Russell, 2017). We then performed a generalized least cally based on the regression information and (2) calcu-
squares Procrustes (GLSP) superimposition while sliding late residuals to see how these values plotted relative to
the semilandmarks to minimize the total bending energy their size. A slope was considered allometric if the prede-
(Gunz & Mitteroecker, 2013; Perez et al., 2006), using the termined isometric slope fell outside the boundaries of
program TpsRelw 1.69 (Rohlf, 2021). Aligned semiland- the confidence intervals. Goodness-of-fit, significance,
marks were averaged for crowns in the same position on and 95% confidence were all reported, with significance
opposite sides. defined as a p-value below 0.05.
The undulating margins of the jaws exists in both Cro- The semilandmarks for both tooth crown shape and
codylia and Spinosauridae, but has never been quantified. jaw shape were examined using a geometric morphometric
Data on jaw margin shape were derived from images of the semilandmark analysis (Bookstein, 1997; Mitteroecker
rostra and dentaries from the lateral perspective. Concern- et al., 2013; Sheets et al., 2004; Zelditch et al., 2004). First,
ing crocodylians, data were collected from both the ros- we conducted multivariate analysis of variance (MANOVA)
trum and dentary from 16 individuals. Only Baryonyx to determine if there were significant differences (p < 0.05)
walkerii and Suchomimus tenerensis had both their rostra between the major taxonomic groups sampled in PAST.
and dentary preserved, so the remaining spinosaurids sam- This was done for the tooth crowns, but could not be done
pled only had one or the other available for study. This for the jaw shape as the sample size was too small. Second,
resulted in six rostra and six dentaries total for spinosaur- a principal component analysis (PCA) was conducted in
ids. Margins of the jaw were traced similar to above using MorphoJ v. 106d (Klingenberg, 2011) to visualize the degree
TpsDig. For both jaws, one curve was drawn from the of shape variance among both teeth and jaws. PCs were
distal-most alveolus to the mesial-most alveolus on the plotted in a series of bi- or tri-variate plots, with vector dia-
tooth-bearing side. The second curve started at mesial-most grams indicating the extremes of shape variance along
alveolus, progressed around the tip of the jaw, and ended each axis.
on the opposite side of distal-most alveolus. This method
considered the tooth-bearing region only, and not any ana-
tomical structures caudal to it. As there are often occlusal 3 | RESULTS
grooves in between the alveoli in crocodylians
(Brochu, 1997), our outline connected the margins of adja- 3.1 | Alveolar area and heterodonty
cent alveolar processes to one another and did not go into
these grooves. The two traced margins were then trans- Spinosaurids, regardless of whether they were baryony-
formed into 60 equidistant coordinates each and the chines or spinosaurines, showed a consistent pattern of
rostral-most landmarks were combined, resulting in three alveolar area along their more mesial positions, yet dif-
discrete landmarks and 117 semilandmarks. This was again fered greatly in their distal positions. All specimens had
followed GLSP superimposition and sliding of the two regions with disproportionately large alveoli on the
semilandmarks. cranial arcade at the positions of premaxillary teeth 2–3
D'AMORE ET AL. 5

F I G U R E 1 Host bones and alveoli for sample members of Spinosaurinae (left) and Baryonychinae (right), including both the fossil and
diagrammatic representations of the alveoli from a single side. Scale bar = 50 mm. (a) Premaxilla and maxilla of MSNM V4047 (photograph
taken by Cristiano Dal Sasso), (b) partial dentary of NHM R16421, (c) premaxilla and maxilla of cast of MNN GDF501, and (d) dentary of
NHM R9951. Silhouettes modified with permission from Scott Hartman. Note the relative differences in the alveolar sizes between the two
major clades.

and maxillary tooth 4, and there was usually a precipi- ‘procumbent’ (Gignac & Erickson, 2014) or a ‘pseudoca-
tous drop in alveolar area between these positions nine’ (Brochu, 1999). The largest premaxillary alveolus
(Figure 1). Spinosaurids all showed a gradual decrease in was at position 4 for all species. Alligatorids had another
alveolar area after maxillary tooth 4 that typically went to large alveolus at maxillary position 4, and crocodylids
the last alveolus, but the arcade itself was eight positions and Tomistoma at position 5. These taxa all had a third
longer in the baryonychine Suchomimus than all spino- set of enlarged alveoli at maxillary positions 9–11 as well,
saurine maxillae (see also Sereno et al., 1998). Concern- which only Gavialis gangeticus lacked. Along the dentary,
ing the dentary, alveoli were enlarged at positions 3–4, all taxa had enlarged alveoli at positions 1 and 4. A third
followed by an immediate drop in size. After these, Bar- enlargement was located at the distal dentary at position
yonyx maintained small alveolar sizes along their remain- 11 for crocodylids, 12–13 for alligatorids and Tomistoma,
ing arcade and had twice as many overall positions as and was again not present in Gavialis (for data, see
spinosaurines (see also Charig & Milner, 1997). Spino- Figure S2 in Appendix S1).
saurines instead showed another region of alveolar When the AAA for each individual was plotted
enlargement along the caudal dentary, with crowns of against both HW and RW, isometric regressions resulted
similar size to the more rostral dentary set. There was a (Figure 2a–c). These indicated larger animals did not nec-
gradual increase in crown size from positions 9 to 12 fol- essarily have relatively larger alveoli. Concerning AAA
lowed by a gradual decrease to the distal-most position. versus RW, spinosaurids showed little overlap with crocody-
Although its taxonomic position is unclear, the incom- lids. The former almost always had larger relative alveoli
plete Iberospinus dentary had a pattern of heterodonty than the latter. This resulted in all but one spinosaurid
most similar to baryonychines (for data, see Figure S1 in maintaining positive residuals, whereas crocodylian resid-
Appendix S1). uals were positive for smaller RW individuals but became
Crocodylians had regions of enlarged alveoli as well, more negative as they got larger. Spinosaurines had consis-
which correlated to the large crowns documented in tently high residuals, but the baryonychines with relatively
numerous prior studies (see D'Amore et al., 2019 and ref- complete jaws (Baryonyx and Suchomimus) had the lowest
erences within). These teeth are often referred to as residuals out of Spinosauridae. Concerning AAA versus
6 D'AMORE ET AL.

F I G U R E 2 Analysis of alveolar metrics regressed against size covariates. Average alveolar area (AAA) for each specimen were regressed
against (a) rosette width (RW) and (b) head width (HW). Morphological disparity (MD) was regressed against (c) AAA, (d) RW, and (e) HW.
All metrics were natural-logarithm scaled, with regression information and statistics indicated. Isometry equals a slope of 2.0 for a–c and 4.0
for d,e. Note the position of the slender-longirostrine crocodylians, which are indicated by hollow data points. For alveolar and covariate
data, see Appendix S1.
D'AMORE ET AL. 7

HW, Suchomimus had the most positive residual and Irrita- plotted against RW, spinosaurids separated from crocody-
tor was in the negative range. All gavialids and Crocodylus lids on the larger end similar to AAA versus RW. Also
niloticus had negative residuals, but the remaining crocody- similar was the distribution of spinosaurine and baryony-
lids were positive with alligators occurring on both sides of chine residuals, although the Baryonyx residual was
the regression. greater.
When size heterodonty (MD) was plotted against any
of the size covariates, significant, positively allometric
regressions resulted (Figure 2d,e). This indicated that 3.2 | Tooth crown morphometrics
larger animals in general had more dental size variation
in their arcades than smaller ones. The MD versus AAA Tooth crown geometric morphometrics yielded only two
regression showed most slender-longirostrine crocody- principal components (PCs) with enough variance to be
lians had relatively low heterodonty for their tooth size considered biologically relevant (Figure 3). Although
based on negative residuals (except for a single Mecistops there was overlap between all theropod groups, the
cataphractus). With the exception of one incomplete ranges of morphotypes within these groups varied. Most
maxilla (CPI 477), spinosaurid specimens of similar sizes spinosaurids had relatively narrow teeth found mostly
had greater residuals than these crocodylians, and over- between 0.05 and 0.20 of PC1. Concerning PC2, spino-
lapped more so with the generalists/macro-generalist spe- saurines had teeth limited to being straight to slightly
cies. Iberospinus had the highest residuals out of all taxa curved. Baryonychines overlapped with them, but also
sampled. The upper extreme of the regression was domi- had a minority of crowns that show more curvature. This
nated by spinosaurines and Crocodylus porosus, both of resulted in them also overlapping with Megalosauridae
which had primarily negative residuals. When MD was as well, which had exclusively curved teeth they were
regressed against HW, both Irritator and Suchomimus only found on the negative side of PC2. When crocody-
had high residuals relative to crocodylians. When MD lians were compared with the theropods, there was

F I G U R E 3 Principal components analysis for tooth crown shape for Crocodylia, Megalosauridae, and Spinosauidae depicting the first
two principal components. Extremes of tooth crown shape variance along each axis are indicated by outlines. Photographs of a range of
tooth morphotypes found within each taxonomic group are indicated, and megalosaurid tooth photographs were contributed by Colin
Boisvert and Christophe Hendrickx. Note the narrow ranges of spinosaurid crown shapes relative to the other taxa. For PC data, see
Appendix S2.
8 D'AMORE ET AL.

substantial overlap but also obvious differences in ranges

of morphology. All three major clades of crocodylians
had a wider range of dental morphotypes along PC1. Cro-
codylians had numerous crowns much shorter and wider
than any spinosaurids, and these accounted for about
half of the shape variance seen in the sample. Spino-
saurids fit entirely within the breadth of the gavialid
PC1 range. Spinosaurid teeth had similar values to
Tomistoma schlegii, but Gavialis gangeticus teeth were
usually more slender/elongate (indicated by lower PC1
values). A substantial portion of crocodylid crowns
with lower PC1 values overlapped with the spinosaur-
ids, many of which fell within the slender-longirostrine
category. Alligatorids overlapped spinosaurids very lit-
tle. Crocodylian crowns are known for having little
curvature, and their relatively positive PC2 values
reflected this. All these taxonomic groups were signifi-
cantly different according to the MANOVA (Wilk's
Λ = 0.0168, F = 3.623, and p < 0.0001).

3.3 | Jaw morphometrics

When geometric morphometric analyses were executed

for the rostrum and dentary, taxa clustered together into
distinct morphotypes for both. For the rostrum, three
PCs were considered biologically meaningful (Figure 4a).
F I G U R E 4 Principal components analysis for rostrum (a) and
All spinosaurids had deep rostra that they shared with
dentary (b) shape for Crocodylia and Spinosauridae representing
generalist/macro-generalist crocodylian species, which the first three principal components. Extremes of the rostrum/
distanced them from the slender-longirostrine crocody- dentary shape variance along each axis are indicated by outlines.
lians along PC1. All spinosaurids also had their premaxil- For PC data, see Appendix S2.
lary convexity at the rostrum tip, giving them a
downturned rosette that crocodylians lacked. In spinosaur-
ines the rosette dipped lower than the remainder of the ros- Spinosaurids plotted in the extremes concerning PC3; Bar-
trum, whereas in baryonychines it is equal with the yonychines displayed upturned rostral dentaries, spinosaur-
maxillary convexity (as observed by Lacerda et al., 2022). ines instead had an enlarged caudal region, and the
Spinosaurines and baryonychines also differ in the position crocodylians were found in between.
of their maxillary convexity: spinosaurines had this convex-
ity very caudally positioned resulting in a lengthy concavity
between their two convexities, yet baryonychines positioned 4 | DISCUSSION
it very rostral resulting in a small concavity. Crocodylians
had neither of these extremes. 4.1 | Baryonychinae versus
Three principal components were considered biologi- Spinosaurinae
cally meaningful for the dentary as well (Figure 4b). As
with the rostrum, spinosaurid dentaries plotted on the This study quantified clear similarities and differences in
deeper end of the spectrum with the generalist/macro- the craniodental apparatus between baryonychines and
generalist crocodylians, although the spinosaurines spinosaurines. Concerning the similarities, all spinosaurids
were more extreme than baryonychines. Spinosaurids had several sets of enlarged alveoli located at the rostral end
were separated from crocodylians on account of an of their jaws (Figure 5). All crocodylians have similarly posi-
enlarged rostral region of the dentary, with slender- tioned enlarged crowns as well (D'Amore et al., 2019; Kieser
longirostrine crocodylians having a more consistent et al., 1993), a trait also shared by ‘front-fanged’ piscivorous
dentary depth overall and generalist/macro-generalist cro- fish (Mihalitsis & Bellwood, 2019). These crowns are typi-
codylians having disproportionately large caudal dentaries. cally apically sharp, and optimally positioned to secure
D'AMORE ET AL. 9

prey. Not only is this region most likely to reach prey first
during an attack, but it will also move fastest as it is farthest
from the jaw joint (Busbey, 1989; Iordansky, 1973). We sus-
pect spinosaurids incorporated a similar strategy, as all spi-
nosaurid crowns sampled here tended to have a relatively
slender, acute morphology ideal for puncturing items
(Figure 6). This morphology, coupled with rapid biting as
indicated by skull shape (Sakamoto, 2010) and quadrate
morphology (Hendrickx et al., 2016), strongly suggests spi-
nosaurids acquired their prey with quick strikes. The sec-
ondary palate reinforced the rostrum to withstand impacts
during these strikes (Busbey, 1994; Rayfield et al., 2007),
and, as both baryonychine and spinosaurine skulls had a F I G U R E 5 Diagrams of a baryonychine (a) and spinosaurine
similar resistance to bending and torsion (Cuff & (b), with differences between the taxa labeled. Labeled features
Rayfield, 2013), this behavior was probably consistent include (1) the length of the rostral concavity, (2) the position and
between the groups. number of enlarged teeth, (3) the size and curvature of the rostral
One of the most obvious differences between the two dentary, and (4) the size and number of additional tooth positions
major spinosaurid clades was that baryonychines possessed in Baryonychinae. Crown size is based on alveolar lengths. Crown
shape is identical across the arcade, representing the average tooth
many more alveoli (Figure 5). The consistent yet smaller
shape for each taxon sampled here.
sizes of these alveoli were what gave the baryonychines that
preserved them lower average tooth sizes. Modern crocody-
lians have variable tooth counts as well (Brown et al., 2015). terrestrial vertebrates [such as a juvenile iguanodon-
Gavialis not only has the most tooth positions by far but also tian (Charig & Milner, 1997)]. These factors together
the smallest relative teeth. Baryonychines and Gavialis also support that the baryonychine diet was not restricted
lacked a region of enlarged distal dentition and had a greater to piscivory or especially small prey. Indeed, spinosaur-
degree of crown curvature than the rest of their respective ids as a whole may have had a broad diet with evidence
clades. Gavialis is known for its diet consisting of mostly of the consumption of fish, terrestrial dinosaurs, and
small aquatic prey dominated by fish (Groombridge, 1982; pterosaurs (Buffetaut et al., 2004; Charig & Milner,
Thorbjarnarson, 1999; Whitaker & Basu, 1982), which it 1997; Hone & Holtz, 2017).
swallows with little processing. Baryonychines were also Another key difference between spinosaurid taxa is
probably doing less extensive processing than their spino- that Spinosaurinae had an additional set of enlarged alve-
saurine relatives, as their distal crowns were probably used oli along their distal dentary that Baryonychinae lacked
primarily for gripping their food during inertial feeding (Figure 5). This condition suggests that their prey
(Figure 6). Although it may be compelling to argue that bar- required a degree of processing before being swallowed
yonychines were fish specialists because of these similarities (Figure 6). All extant crocodylians (except Gavialis)
to Gavialis, we caution against this for several reasons. have enlarged distal dentary and distal maxillary
(1) Most Gavialis teeth were thinner and more elongate than crowns (Figure S2), and their morphology usually cor-
any spinosaurid teeth in our sample, indicating they had relates to the durability of their prey type (Aoki, 1989).
lower bending strengths and were less resistant to prey Crocodylians propel food to this region of the jaw using
forces. Spinosaurid teeth were more similar to slender- inertial feeding (Cleuren & De Vree, 2000; Davenport
longirostrine crocodylids/Tomistoma, and there are et al., 1990), and being closer to the hinge allows these
numerous reports of these crocodylians eating hard, teeth to impact the food with more force for puncturing
large, and/or terrestrial prey (Brazaitis, 1973; and/or crushing (Cleuren et al., 1995;Erickson
Galdikas & Yeager, 1984; Groombridge, 1982; et al., 2003, 2012; McHenry et al., 2006). Their distal
Selvaraj, 2012). (2) Baryonychine alveolar sizes were teeth are also relatively more molariform, increasing
both relatively and absolutely greater than Gavialis bending strength and making them more resistant to
(as well as much of Crocodylia). (3) Baryonychines breakage (Gignac & Erickson, 2014; Monfroy, 2017;
were more heterodont than most slender-longirostrine Valkenburgh & Ruff, 1987). Our data shows that the
crocodylians, especially Gavialis. (4) Gavialis has a rel- most robust crowns exist in generalist/macro-
atively weak skull (Walmsley et al., 2013), with Baryo- generalist alligatoroids and crocodylids, and these are
nyx outperforming it concerning bending and torsion often used to break down hard prey items such as mol-
(Cuff & Rayfield, 2013). (5) Putative stomach contents lusks, crustaceans, tetrapod bones, and even turtle
suggest that Baryoynx ate, at least on occasion, small shells (Erickson et al., 2003; Gignac & Erickson, 2014).
10 D'AMORE ET AL.

F I G U R E 6 Feeding behavior in a sample baryonychine (top) and spinosaurine (bottom), including before (a), during (b), and after (c) a
strike. The starred images indicate behaviors resulting in deep punctures. Note the relative prey size and number of starred behaviors.

Spinosaurine crowns instead overlap with Tomistoma limited to aquatic prey that were small for their size.
and slender-longirostrine crocodylids, which are not Slender-longirostrine crocodylians tend to consume prey
particularly durophagous (Erickson et al., 2012; much smaller than their body weight (Drumheller &
Peyer, 1968; Webb et al., 1982). Spinosaurid bite force Wilberg, 2020), but spinosaurids of similar size had teeth
may have been relatively low for their size range that were both larger and more heterodont. Large crocody-
(Schade et al., 2023), but it was still substantial. In par- lians with high size heterodonty [such as Crocodylus poro-
ticular, the bite force of Spinosaurus was similar to sus (D'Amore et al., 2019)] tend to prioritize large
medium-sized tyrannosaurids (Sakamoto, 2010). This vertebrates (Hanson et al., 2015; Kar & Bustard, 1983;
altogether suggests that the spinosaurines were well Taylor, 1979), and it is possible large spinosaurids did
suited for pulverizing/puncturing prey that could have the same.
been relatively large, but not relatively hard, such as Both spinosaurids and crocodylians independently
gars and Onchopristis (Beevor et al., 2021). evolved size heterodonty with several isolated regions of
enlarged teeth and small teeth in between. This begs the
question: what is the advantage of this morphotype? It is
4.2 | Heterodonty, rostrum shape, and certainly not the only dental morphotype possessed by
prey characteristics aquatic feeding amniotes. Many anisodont plesiosauro-
morphs had crown size changes that occurred in a grad-
Overlapping in heterodonty with generalist/macro-general- ual, as opposed to oscillating, fashion (Kear et al., 2017;
ist crocodylians also suggests that spinosaurids were not Sassoon et al., 2015). Other aquatic predators simply
D'AMORE ET AL. 11

evolved low heterodonty, such as teleosaurid crocodylo- accommodate this depth. CT scans of spinosaurid jaws
morphs (Johnson et al., 2022) and odontocete whales indicate that much of the host bone volume was taken up
(Rommel, 1990). The answer may lie in the necessity to by roots (Rayfield et al., 2007). Lacerda et al. (2022) found
create substantial punctures in prey, as opposed to simply that premaxillary morphology was strongly linked with
gripping it. If a single tooth on one of the rows were to allometry, with more downturned rosettes in larger taxa.
initially contact prey during a strike, then the maximum As we have determined that heterodonty is also allome-
amount of stress will be delivered at its apex tric, it seems plausible that the greater differences in rela-
(Anderson, 2018; Mihalitsis & Bellwood, 2019). This tive tooth sizes may have generated this trend. It has also
would maximize the likelihood of substrate failing and been suggested that brachyrostral phytosaurs were, in
the tooth puncturing. Alternatively, similarly-sized, adja- certain aspects, better mechanical/ecological analogues
cent crowns would instead simultaneously contact prey for spinosaurids as they share this downturned rosette
during a strike. The stress delivered would consequently (Yun, 2024). This very well may have been the case, but
be divided between their multiple apices, resulting in this convergence may have been driven by the fact that
shallower punctures or none at all. The morphotype seen both taxa have large/small teeth in similar positions. Dis-
in the spinosaurid jaw could have been crucial in secur- tantly related eureptiles associated with a variety of envi-
ing large, tough skinned, and/or struggling prey, as the ronments, such as captorhinids, dilophosaurids, and
bite force would be focused on the few enlarged crowns. proterosuchids, also reflect this trend (Abel et al., 2022;
Conversely, having adjacent crowns of consistent size Marsh & Rowe, 2020). Future anatomical studies should
would not have been a problem if the prey was small or consider this constraint as a potential driver of jaw
highly compliant (Mihalitsis & Bellwood, 2019). For architecture.
example, odontocete whales hunt prey that is usually Even if jaw margin undulations are developmentally
under 10% of their body length (MacLeod et al., 2006, constrained by alveolar depth, this does not preclude
2007), so there is little need to maximize tooth puncture them from having functional importance when feeding.
depth. The concavities of one jaw may have interlocked with the
The pattern of large and small teeth in both Spino- convexities of the other. Most reconstructions position
sauridae and Crocodylia coincides with the undulating the rostral-most dentary within the rostral-most concav-
convexities and concavities along their jaw margin. These ity of the upper jaw during occlusion (Ibrahim
two phenomena appear to be coupled; large teeth are et al., 2020; Sereno et al., 2022). This region seems ideal
always located on the convexities, and small teeth to pin prey in, secured in place by the enlarged dentary
are found in the concavities. As different sized teeth teeth. Interlocking jaw margins also occur in many of the
change location along the jaw, so do these corresponding crocodylians sampled here, as well as the taxonomically
convexities and concavities. Both baryonychines and spi- distant Pike Conger eel (Vullo et al., 2016), presumably
nosaurines share enlarged crowns at maxillary 4, but, in for similar purposes. In baryonychines this upper jaw
order to accommodate their additional distal teeth, baryo- concavity was very rostrally positioned and relatively
nychines moved this position rostrally. The correspond- small, and the dentary was curved so to fit into
ing maxillary convexity moved with it, shortening the it. Spinosaurines, on the other hand, had a very long con-
concavity between it and the premaxillary convexity cavity that interlocked with their bulbous, uncurved den-
(Figure 5). Additional dental enlargements resulted in tary. Even if both groups used this structure in a similar
additional changes in jaw depth, as the spinosaurine spe- manner, it most certainly would influence the relative
cific hypertrophied distal teeth were coupled with a dee- size of the prey that could be positioned there. This lends
per caudal dentary. The same phenomenon can be seen further support to the idea that the spinosaurine jaw was
across Crocodylia, with more size heterodont a crocody- better adapted to deal with larger prey than baryonchines
lians having more dramatic the undulations. The down- (Figure 6). In addition to the undulating margin, spino-
turned rosette existing in spinosaurids but not saurids also laterally compressed their jaws. This may
crocodylids appears to be coupled with the fact that the have been an adaptation to reduce drag when striking
former has their largest premaxillary alveoli at positions prey within water. The rostrum was especially com-
2 and 3 as opposed to 4 (D'Amore et al., 2019). pressed at the rostral concavity, suggesting that this fit in
The coupling of tooth size and jaw undulation between the enlarged dentary teeth during jaw closure.
strongly suggests this may be a developmental constraint. The lateral expansion of the rosette may therefore repre-
Large teeth typically have long roots, and alveoli need to sent a trade-off; it may have been widened in order to
be deep enough to fit them as well as their developing increase the chances of contacting prey during a strike,
replacement teeth. The convexities on the jaw margin but its maximum width may have been limited due to the
may simply exist to allow for the space needed to drag it created.
12 D'AMORE ET AL.

4.3 | Dental function and interaction often consume prey larger than their mouths, and use a
with prey ‘Death Roll’ to disarticulate it into manageable portions
(Drumheller et al., 2019; Fish et al., 2007). Although it is
Spinosaurid dentition reflects a derived morphotype dis- highly unlikely that spinosaurids rolled, they may have
tinct from other theropods. Most theropods, including instead used their enlarged forelimbs and claws to reduce
the megalosaurids sampled here, had ziphodont dentition prey. The function of the forelimbs has been speculated
with distally curved apices, denticulate carinae, and a upon with little consensus [see Hone & Holtz, 2017 for a
degree of lateral compression (Hendrickx et al., 2019). synopsis] as they could flex and extend powerfully, facili-
Concerning the crown outline, the spinosaurid morpho- tated by an enlarged deltopectoral crest and olecranon
type is rather constrained as most teeth were elongate (Charig & Milner, 1997; Ibrahim et al., 2014; Sereno
and narrow. Although not directly measured, lateral et al., 1998). The manus could most likely reach the jaw
compression and denticulation can be quite variable and and food secured in it. The possibility therefore existed
sometimes not present (Hone et al., 2010; Kellner & that the claws assisted in manipulating the food, breaking
Campos, 1996). One could subsequently ask what func- up large prey that could neither be easily swallowed
tional changes resulted from this morphological transi- whole nor could be reduced by the jaws alone given their
tion away from ziphodonty. Theropod tooth marks and lack of cutting action. Buoyant food would not have been
biomechanics often suggest puncture-and-pull behavior easily secured with the feet as theorized for land thero-
(Erickson & Olson, 1996; Hone & Watabe, 2010; pods (Fowler et al., 2011), so this manus/jaw combina-
Snively & Russell, 2007; Snively et al., 2013; Torices tion would be particularly useful when feeding in water.
et al., 2018): their crowns would enter substrate and be
drawn in the distal direction. The curved apices allowed
for axial loading in this direction, lateral compression 4.4 | Other considerations
reduced resistance from substrate, and the denticles facil-
itated cutting of flesh (Frazzetta, 1988). This behavior is Positive allometry positions the largest spinosaurids, such
seen in the Komodo monitor (Varanus komodoensis), as Spinosaurus, as highly heterodont based on their size
which has distinct ziphodont dentition alone. In contrast the much smaller Irritator had rela-
(Auffenberg, 1981; D'Amore & Blumenschine, 2009). As tively small alveoli, contributing to its large inter-dental
spinosaurid, and especially spinosaurine, crowns became spacing observed by Schade et al. (2023). Similar allome-
straighter, the tooth would have to move entirely in a tric changes in Crocodylia have been well documented
downward motion to facilitate axial loading. As the teeth and are often seen as facilitating ontogenetic niche shifts
would no longer be drawn distally, there would be no (Dodson, 1975; Erickson et al., 2003; Gignac &
need for compression or denticles. The crowns could now Erickson, 2016; Gignac & O'Brien, 2016). Numerous stud-
be thickened labio-lingially, increasing bending strength ies have suggested that major shifts in trophic niche
about this axis. In summary, spinosaurid crowns occur during theropod ontogeny as well (Holtz, 2021;
increased their ability to puncture and resist lateral forces Johnson-Ransom et al., 2023; Lockley & Xing, 2021;
at the expense of cutting through substrate (Holtz, 1998). Rowe & Snively, 2022; Schroeder et al., 2021). There are
Modern crocodylian teeth also do not function to cut like currently few fossils of juvenile spinosaurids described
ziphodont teeth. They primarily puncture and secure though (Maganuco & Dal Sasso, 2018), but these were
prey, which is especially important if the prey is to be found in semiaquatic habitats and shared the rosette
drowned, moved, or disarticulated. Heckeberg and Rauhut morphology with adults (Lakin & Longrich, 2019). Cau's
(2020) suggest rapid tooth replacement was a specialization (2024) recent phylogenetic study returned Scipionyx sam-
for having teeth that functioned in this manner, as the niticus as potentially a very immature spinosaurid.
forces exerted by struggling prey would increase the risk of Although lacking the derived traits typical of the clade,
losing teeth during feeding. A fast replacement rate kept its gut contents included both terrestrial prey and fish
the tooth row functional under these circumstances. (Dal Sasso & Maganuco, 2011). This tentatively suggests
Reducing their ability to cut flesh points to a funda- that young spinosaurids shared feeding habitats with
mental shift in food processing for spinosaurids. It may adults.
have made them less effective at scavenging from large Our detailing of the spinosaurid feeding apparatus
carcasses like those of adult sauropods, where removing does not provide strong validation for either subaqueous
sections of meat would have been comparatively easy for pursuit predator or the wading feeding hypotheses. Most
most theropods (Lei et al., 2023). This does not necessar- raptorial marine amniotes have reduced heterodonty
ily mean that prey sizes were limited to objects that (Johnson et al., 2022; Rommel, 1990), but the crocody-
needed to be swallowed whole though. Crocodylians lians which were most similar in heterodonty to
D'AMORE ET AL. 13

spinosaurids feed both subaqueously and on terrestrial opportunistic, and will consume most available food items
prey from the water's edge (Galdikas & Yeager, 1984; they deem worthwhile (Grigg & Kirshner, 2015). Even
Selvaraj, 2012). As no crocodylian has been observed though high size heterodonty in crocodylians is usually
feeding by bringing its jaw down into the water, they are linked to larger prey sizes, there are C. niloticus populations
overall not a very good model organism for the wading or reported to be mostly, and sometimes exclusively, piscivo-
shoreline feeding hypotheses. In summary, our data does rous (Cott, 1961; Graham, 1968). Spinosaurids were likely
not strongly favor one hypothesis over the other. similarly opportunistic, which makes local environmental
There are several caveats and unknown factors to be circumstances, such as habitat range(s) and type(s), prey
considered when drawing inferences from these results. availability, and seasonal changes, as important in recon-
For a detailed synopsis see Appendix S1. structing niche occupation as morphology. This reality
should caution us against assuming that unique morpho-
types like the spinosaurid feeding apparatus are inherent
5 | C ON C L U S I ON indicators of a narrow niches or prey specialization. Traits
such as these may have indeed been selected to handle a
Through multiple avenues of data collection and analysis, specific food type, but that does not mean that said food
we were able quantify aspects of the craniodental apparatus makes up all, or even the majority, of the animal's diet
in both Spinosauridae and modern Crocodylia. Spinosaur- [such as hard food adaptations in varanids (D'Amore, 2015)
ids had a mosaic of craniodental characters, some of which and australopiths (Strait et al., 2013)]. Future fossil discover-
were similar to crocodylians and others that were quite dif- ies and the application of novel techniques should result in
ferent. This calls into question the idea that they were croc- a clearer perspective on the life history of these unique and
odile “mimics,” but still leaves room for interpretation of fascinating theropods.
spinosaurid paleoecology by way of analogy. To summarize,
spinosaurids had tooth crowns similar to certain slender- AUTHOR CONTRIBUTIONS
longirostrine crocodylians in shape but heterodonty and Domenic C. D'Amore: Writing – original draft; conceptu-
rostral morphology more similar to generalists/macro-gen- alization; investigation; methodology; visualization;
eralists. Dentition was also absolutely large compared to writing – review and editing; formal analysis; validation;
crocodylians. This indicates the spinosaurid feeding appara- data curation; resources. Evan Johnson-Ransom:
tus was well suited for quickly striking, puncturing, and Writing – review and editing; writing – original draft;
holding relatively soft prey, but was most likely not limited investigation; resources; methodology. Eric Snively:
to fish or small prey. Spinosaurid dentition allowed for Writing – original draft; writing – review and editing; con-
puncturing at the expense of cutting flesh or durophagy. ceptualization; validation. David W. E. Hone: Resources;
Size heterodonty was high due to isolated sets of enlarged conceptualization; writing – original draft; writing – review
teeth, which applied maximum force to prey for deep punc- and editing; validation; methodology; investigation.
tures. This was coupled with an undulating jaw margin that
allowed for spinosaurids to pin and hold prey, which could ACKNOWLEDGMENTS
have then been subdued, transported, reduced with the We would like to thank the curators and collections staff
forelimbs, and/or pushed deeper into the mouth. Baryony- at the American Museum of Natural History, Natural
chines probably focused on prey items that needed little History Museum, and Royal Ontario Museum for allow-
oral processing as indicated by numerous, small distal teeth, ing us access to their collections. Paul Sereno also
whereas spinosaurines had a suite of traits indicating they allowed us access to his professional collections. Colin
could have hunted and processed relatively large prey. Boisvert, Cristiano Dal Sasso, Christophe Hendrickx,
Lengthier convexities/concavities in jaw margin and bigger Matthew Lamanna, and Simone Maganuco contributed
teeth were for holding larger objects in place, and hypertro- photographs of fossils. Scott Hartman allowed us access
phied distal crowns allowed for more extensive oral proces- to his reconstructions. This research was partially funded
sing. In particular, Spinosaurus had high size heterodonty by the Daemen University Faculty Research Award. We
resulting from allometric scaling. are grateful for all these sources.
A goal of much spinosaurid research is to reconstruct
the ecology of an unusual extinct clade. Our morphologi- C O N F L I C T O F I N T E R E S T S T A TE M E N T
cal analysis, combined with comparisons with modern The authors declare no conflicts of interest.
crocodylian analogues, sheds light on the fundamental
niches of spinosaurids and associated behaviors. Determin- ORCID
ing the actual prey consumed, or realized niche, is much Domenic C. D'Amore [Link]
more elusive. Large modern reptilian predators are typically 4376
14 D'AMORE ET AL.

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