# First report of neosuchian remains in Aptian–Albian marine deposits of southeastern France

Jeremy E Martin, Romain Jattiot

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Jeremy E Martin, Romain Jattiot. First report of neosuchian remains in Aptian–Albian marine deposits of southeastern France. Proceedings of the Geologists’ Association, 2025, 136 (5), pp.101132. 10.1016/j.pgeola.2025.101132. hal-05285621

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# First report of neosuchian remains in Aptian–Albian marine deposits of southeastern France

<figure><img src="image_2.png" /></figure>

Jeremy E. Martin<sup>a,*</sup>, Romain Jattiot<sup>b</sup>

## ARTICLE INFO

**Article history:**

Received 18 April 2025

Received in revised form 4 July 2025

Accepted 4 July 2025

Available online 23 July 2025

**Keywords:**

Crocodylomorpha

Neosuchia

Aptian

Albian

Europe

## ABSTRACT

The European fossil record of crocodylomorphs during the Cretaceous is indubitably skewed towards the lowermost and uppermost strata. Neosuchian remains from the Aptian–Albian interval are exceptionally rare in marine deposits globally. Yet, they potentially hold important information for understanding the phylogenetic interrelationships of several neosuchian lineages. Here, we assign a single squamosal to cf. Pholidosauridae and some postcranial elements to Neosuchia indet. from Aptian and Albian marine deposits in southeastern France. Although it is presently impossible to ascertain the continental *versus* marine origin of the specimens, their rare occurrence calls for future investigations of the diversity and interrelationships of neosuchians during the mid-Cretaceous.

© 2025 The Author(s). Published by Elsevier Ltd on behalf of The Geologists' Association. This is an open access article under the CC BY license (http://creativecommons.org/licenses/by/4.0/).

## 1. Introduction

Despite their presence in marine ecosystems of the Early and Late Cretaceous (e.g., Chiarenza et al., 2015; Jouve et al., 2021), marine crocodylomorphs (Thalattosuchia, Pholidosauridae or Dyrosauridae) are unknown in marine deposits of the Hauterivian–Albian interval (Martin et al., 2014a). A possible exception is a single jaw fragment from the Aptian–Albian of the Isle of Wight, UK (Young et al., 2014) but whether this specimen was reworked from the Kimmeridge Clay Formation, lived or was washed into the marine environment, is unknown. In parallel to marine ecosystems, European continental ecosystems of the Cretaceous have yielded neosuchian remains at different intervals, notably during the Berriasian–Barremian (e.g., Buscalioni and Sanz, 1990; Salisbury et al., 1999; Martin et al., 2016, 2020) but Aptian–Albian discoveries are represented by too few records (Buscalioni et al., 2011, 2013). Small exquisite specimens referred as *Pietraroiasuchus ormezzanoi* from the Albian of Italy (Buscalioni et al., 2011) occur in a paralic environment, highlighting that freshwater ecosystems did represent a potential source of carcasses into marine deposits. The imperfect Aptian–Albian fossil record of crocodylomorphs prevents testing phylogenetic hypotheses on the interrelationships of several lineages that went extinct or radiated during the mid-Cretaceous.

Of potential relevance for filling this gap in the fossil record, marine deposits of the Aptian–Albian of the Vocontian Basin in southeastern France occasionally yield allochthonous remains of continental origin. Such rare records account for the proximity of emerged lands as indicated by marine sequences that produced amber (Onoratini et al., 2009; Saint Martin et al., 2021) as well as much rarer examples of dinosaur remains (Gervais, 1848–1852; Accarie et al., 1995). However, crocodylomorphs were never reported despite their usual abundance in freshwater ecosystems. In this context, continental vertebrates are underrepresented and enter the marine environment as floating carcasses, explaining their rarity when marine vertebrates are regionally represented by much more complete skeletons (e.g., Vincent et al., 2020). Here, we report for the first time on new neosuchian specimens from the Aptian–Albian interval, highlighting that intensive field survey efforts will probably yield important evidence for filling a gap in the neosuchian fossil record and for assessing their auto- *versus* allochtony.

Institutional abbreviations: UCBL-FSL, Université Claude Bernard Lyon 1; UM-CNL, Université de Montpellier, Carniol locality.

## 2. Geological setting

The Salazac and Carniol sites are located in the southeastern France sedimentary basin (Fig. 1). During the Albian, the central portion of the basin is represented by a pelagic facies (documented from the neighbouring Mont Risou succession; Gale et al., 1996, 2011) and constitutes the Vocontian trough, whereas the shallow

<footnote><sup>a</sup> Université Claude Bernard Lyon1, LGL-TPE, UMR 5276, CNRS, ENSL, UJM, F-69622 Villeurbanne, France</footnote>

<footnote><sup>b</sup> CR2P – Centre de Recherche en Paléontologie – Paris MNHN, Sorbonne Université, CNRS 8 rue Buffon F-75005, Paris, France</footnote>

<footnote>* Corresponding author.
E-mail address: jeremy.martin@cnrs.fr (J.E. Martin).</footnote>
<figure><img src="image_3.png" /><figcaption>**Fig. 1.** Palaeogeographical context of Western Europe during the A, Aptian and B, Albian. The approximative depositional areas of the crocodylomorph specimens described in this study are indicated by red circles.

Maps generated by Deep Time Maps™ 2020, Ron Blakey.</figcaption></figure>

peripheral domains are characterised by glauconitic sands (Médioni, 1984; Bréhéret, 1997; Ferry, 1999; Amédro, 2008). In between are recorded hemipelagic facies (mainly sandy marls), documented from the neighbouring Marcoule succession (Amédro, 2008). The Salazac locality represents a shallow peripheral domain, with a sedimentary record predominantly consisting of glauconitic sands. The material from Salazac described herein was retrieved from a ~0.7 m thick transgressive, condensed level of glauconitic and phosphatised sands (see Jattiot et al., 2021 for further details). The Salazac condensed level represents the 'lower Vraconnian' (Breistroffer, 1936, 1940a, 1940b; Amédro, 2008) with *Mortoniceras* (M.) *fallax* (Breistroffer, 1940a) and *Neophlyticceras* (N.) *blancheti* (Pictet and Campiche, 1858-64), corresponding to the Mortoniceras fallax Zone in the standard zonation (Reboulet et al., 2018; Jattiot et al., 2021) and to UA 13 in the recently published late Albian quantitative biochronology (Jattiot et al., 2023). The material from Salazac was discovered by one of us (RJ) during fieldwork concerned with taxonomic reappraisal of the latest Albian (*Mortoniceras fallax* Zone) Salazac cephalopod fauna (see Jattiot et al., 2021). The Carniol locality was briefly mentioned in Moullade et al. (2006) and is late Aptian in age. Compared to Salazac, the upper Aptian sediment record at the Carniol locality consists of deepwater marlstones (Ferry et al., 2022).

### 3. Systematic palaeontology

* Crocodylomorpha Hay, 1930

* Mesoeucrocodylia Whetstone and Whybrow, 1983

* Neosuchia Benton and Clark, 1988

* cf. Pholidosauridae Zittel and Eastman, 1902

**Age and locality:** late Aptian of Carniol, Vaucluse department, France.

**Referred material:** UM-CNL-1, a right squamosal.

**Description.** The specimen consists of an isolated right squamosal (Fig. 2). The dorsal surface is ornamented with circular to ovoid deep pits in the range of 3 to 5 mm in diameter (Fig. 2A). The distal process that contacted the postorbital is broken off, as well as the posterodistal process that descended on the quadrate. The medial process is nearly complete and preserves the sutural area for the parietal. The anteromedial margin of the bone preserves a wide concave and smooth border that corresponds to the posterolateral margin of the supratemporal fenestra. The medial process presents unparalleled anterior and posterior margins, giving to it a medially tapering profile. The medial process of the squamosal

is about as thick as the anterior process. Given these morphological features, the supratemporal fenestra was positioned close to the posterior margin of the skull table. In ventral view, the sutures for the exoccipital and quadrate can be identified (Fig. 2B) with the dorsal roof for the otic area being placed just lateral to the latter. In lateral view, the posterior lip for the attachment of the earflap musculature is also visible (Fig. 2D).

*Comparisons.* Anatomical comparisons were made from the literature using the descriptions of the following Cretaceous taxa listed in Table 1. Although fragmentary, the squamosal indicates an individual with a relatively small body length, perhaps not surpassing a metre and resembling the pholidosaurid *Terminonaris robusta*. Comparisons were made with several taxa representing different neosuchian lineages (Fig. 2F-L and Table 1). The absence of a ridge on the dorsal surface of the squamosal negates a referral to *Paralligator*, *Wannchampsus* or *Varanosuchus*. The lateral margin of the squamosal, above the otic notch, is slightly convex, as is the case in several neosuchians such as *Bernissartia fagesii*, *Terminonaris robusta* and *Hylaeochampsa vectiana*, but unlike the condition in *Isisfordia duncani*, *Rugosuchus*, *Pholidosaurus* or *Goniopholis* where this area is concave. The medial process of the squamosal is more elongated and proportionally much thinner than that of *Acynodon iberoccitanus*, *Isisfordia duncani*, *Bernissartia fagesii*, *Goniopholis simus*, *Hylaeochampsa vectiana*, *Pietraroiasuchus ormezzanoi*, *Koumpiodontosuchus aprosdokiti*, and *Elosuchus cherifiensis*. Unlike the taxa cited above, the posterior margin of the supratemporal fenestra in the Carniol specimen is close to the posterior margin of the skull table. Its converging margins are more reminiscent of *Rugosuchus nonganensis* and *Terminonaris robusta*. This is also in agreement with wider than long supratemporal fenestrae. Although the squamosals of *Portugalosuchus azenhae* are mostly missing, their margins bordering the supratemporal fenestra differ from that of the Carniol specimen in possessing a wide smooth dorsal surface just anterior to the ornamented bone. Ornamentation patterns devoid of marked pits are characteristics of dyrosaurids but also of *Hylaeochampsa*, and do not resemble the morphology of the Carniol specimen.

In conclusion, the relative posterior position of the supratemporal fenestra on the skull table and the tapering medial process of the squamosal are reminiscent of *Rugosuchus nonganensis* and *Terminonaris robusta*, but the lateral outline above the otic notch is unlike that of *R. nonganensis*. Although morphological evidence is extremely limited, the squamosal of the Carniol specimen resembles that of a longirostrine neosuchian such as the pholidosaurid *Terminonaris robusta*.
<figure><img src="image_4.png" /><figcaption>**Fig. 2.** The right squamosal of an indeterminate neosuchian from the Aptian of Carniol, France in A, dorsal; B, ventral; C, occipital and D, right lateral views. E, Hypothetical reconstruction of the dorsal outline of the Carniol neosuchian as to show the position of the squamosal. The squamosal was mirrored to generate the left counterpart. Comparison of the dorsal aspect of the Carniol squamosal (F) with those of contemporaneous neosuchians. G, *Terminonaris robusta* (redrawn from Wu et al., 2001a); H, *Rugosuchus nonganensis* (redrawn from Wu et al., 2001b); I, *Isisfordia duncani* (redrawn from Salisbury et al., 2006); J, *Bernissartia fagesii* (redrawn from Martin et al., 2020); K, *Hylaeeochampsa vectiana* (redrawn from Clark and Norell, 1992); L, *Pholidosaurus* sp. (redrawn from Martin et al., 2016). Abbreviations: dps, distal process of the squamosal; exos, exoccipital suture; mps, medial process of the squamosal; on, otic notch; pdp, posterodistal process; ps, parietal suture; STF, supratemporal fenestra.</figcaption></figure>

Neosuchia indet.

Age and locality: late Albian of Salazac, Gard department, France.

Referred material: UCBL-FSL 532464, a set of two pathologic vertebrae and UCBL FSL 532465, a femur.

Descriptions.

*Femur.* A left femur is nearly complete, missing its distal condyle on the medial side (Fig. 3A–F). Although the shaft is fractured transversely

at different levels and some cortical bone fragments along the fractures are missing, the general morphology, muscle scars and insertion areas are clearly identifiable. A set of five longitudinal ridges extend along the medial surface of the distal condyle. Similar ridges are observed on the medioproximal head. The femur has a proximodistal length of 250 mm and a width at mid-shaft of 30 mm (Table 2). At mid-shaft, the section is triangular. The dorsal surface of the femur is twisted

<table><caption>**Table 1**

List of crocodylomorph specimens used for comparison. This list is not exhaustive and is restricted to specimens preserving the squamosal.</caption><thead><tr><th>Family</th><th>Taxon</th><th>Stage and location</th><th>Source</th></tr></thead><tbody><tr><td>Eusuchia indet.</td><td>Portugalosuchus azenhae</td><td>Cenomanian, Portugal</td><td>Mateus et al. (2019)</td></tr><tr><td>Hylaeeochampsidae</td><td>Acynodon iberocctanus</td><td>Campano-Maastrichtian, France</td><td>Martin (2007)</td></tr><tr><td>Hylaeeochampsidae</td><td>Hylaeeochampsa vectiana</td><td>Barremian, UK</td><td>Clark and Norell (1992)</td></tr><tr><td>Hylaeeochampsidae</td><td>Pietraroiasuchus ormezzanoi</td><td>Albian, Italy</td><td>Buscalioni et al. (2011)</td></tr><tr><td>Susisuchidae</td><td>Isisfordia duncani</td><td>Albian, Australia</td><td>Salisbury et al. (2006)</td></tr><tr><td>Susisuchidae</td><td>Susisuchus anatoceps</td><td>Aptian, Brazil</td><td>Salisbury et al. (2003)</td></tr><tr><td>Bernissartiidae</td><td>Bernissartia fagesii</td><td>Barr.-Aptian, Belgium</td><td>Martin et al. (2020)</td></tr><tr><td>Bernissartiidae</td><td>Koumpiodontosuchus aprosdokiti</td><td>Barremian, UK</td><td>Sweetman et al. (2014)</td></tr><tr><td>Paralligatoridae</td><td>Paralligator gradilifrons</td><td>Santonian, Mongolia</td><td>Kuzmin et al. (2025)</td></tr><tr><td>Paralligatoridae</td><td>Wannchampsus kirpachi</td><td>Aptian, Texas</td><td>Adams (2014)</td></tr><tr><td>Paralligatoridae</td><td>Tarsomordeo winkleri</td><td>Aptian, Texas</td><td>Adams (2019)</td></tr><tr><td>Paralligatoridae</td><td>Rugosuchus nonganensis</td><td>Latest E. Cretaceous, China</td><td>Wu et al. (2001b)</td></tr><tr><td>Atoposauridae</td><td>Varanosuchus sakonnakhonensis</td><td>Valan.-Hauter., Thailand</td><td>Pochat-Cottilloux et al. (2024)</td></tr><tr><td>Goniopholididae</td><td>Goniopholis simus</td><td>Berriasian, UK, Germany</td><td>Salisbury et al. (1999)</td></tr><tr><td>Pholidosauridae</td><td>Terminonaris robusta</td><td>Turonian, Montana</td><td>Wu et al. (2001a)</td></tr><tr><td>Pholidosauridae</td><td>Pholidosaurus sp.</td><td>Berriasian, France</td><td>Martin et al. (2016)</td></tr><tr><td>Pholidosauridae</td><td>Elosuchus cherifiensis</td><td>Albo-Cenomanian, Morocco</td><td>de Lapparent de Broin (2002)</td></tr><tr><td>Dyrosauridae</td><td>cf. Rhabdognathus sp.</td><td>Maastr.-Danian, Mali</td><td>Brochu et al. (2002)</td></tr></tbody></table>
<figure><img src="image_5.png" /><figcaption>**Fig. 3.** Postcranial elements of the new neosuchian from the late Albian of Salazac, Gard, France. Left femur in A, proximal; B, distal; C, dorsal; D, medial; E, ventral and F, lateral views. Fused thoracal vertebrae in G, dorsal; H, right lateral; I, ventral; J, anterior and K, posterior views. Abbreviations: cal, callus on suture; dep, depression on proximodorsal surface; k, knob; Ihc, distal lateral hemicondyle; ns, neural spine; poz, post-zygapophysis; prz, pre-zygapophysis; 4t, fourth trochanter; arrows point anteriorly.</figcaption></figure>

mediolaterally. The fourth trochanter is prominent and sits at about 80 mm from the proximal condyle. The most distinctive character of the femur is the presence of a deep and wide depression on the antero-lateral surface of the proximal head (Fig. 3C, D).

**Vertebrae.** Two thoracal amphicoelous vertebrae from the posterior region of the trunk are preserved with their centra being fused to each other (Fig. 3G–K). They are fairly complete but are broken off along their edges including the anteroventral margin of the anterior centrum and lack most of the transverse processes and the distal tips of the neural spines. There are no dia- and parapophyses on the centrum; on the transverse processes, their positions cannot be evaluated. The anterior surface of the centrum of the anteriormost vertebra is concave and the posterior surface of the posterior vertebra is flat with bevelled margins. The centra are about as long as wide and slightly compressed dorsoventrally (35 × 40 × 30 mm), the posterior surface of the centrum being half a centimetre wider than the anterior margin. The first centrum is slightly shorter than the second one (38 versus 35 mm). The fusion between the centra appears to be pathological as evidenced by an irregular and asymmetric thickened area or callus, best visible along their ventral surfaces or at the articulation between both centra (Fig. 3H, I). The right transverse process of the anteriormost vertebra is broken off just lateral to the pre-zygapophysis, showing that the transverse process originates

<table><caption>**Table 2**
Measurements of the new femur from Salazac following Farlow et al. (2005).</caption><tr><td>FL (femur length)</td><td>250 mm</td></tr><tr><td>Fdh (femur distal height)</td><td>36 mm</td></tr><tr><td>Fdw (femur distal width)</td><td>7 mm</td></tr><tr><td>Fpmn (femur minimum proximal diameter)</td><td>23 mm</td></tr><tr><td>Fpmx (femur maximum proximal diameter)</td><td>58 mm</td></tr><tr><td>Ftr (distance from the most proximal point on the proximal articular surface of the bone to the fourth trochanter)</td><td>81 mm</td></tr><tr><td>Fc (minimum midshaft circumference)</td><td>30 mm</td></tr></table>

from the upper margin of the centrum. Its section is dorsoventrally compressed. The prezygapophyses are facing fully dorsally and are horizontal as are the post-zygapophyses. The pre-zygapophyses are large, measuring more than 20 mm in width, and reaching laterally further than the smaller post-zygapophyses. The pre-zygapophyses project as far as the anterior margin of the centrum. The neurocentral suture is fused and not discernible. Laminae and pleurocoele are visible on the right lateral side of the specimen (Fig. 3H). The neural canal is wider than high. The neural spines are vertical on their anterior side and slope posteriorly.

**Comparisons.** The associated vertebrae, despite the presence of a parthology (fusion of the centra), are distinctively amphicoelous; attribution to the Eusuchia can therefore be excluded. The femur has the characteristic sigmoid outline of aquatic or semi-aquatic crocodylomorphs. A marked depression on the anterolateral surface of the proximal head is generally absent in the femora of thalattosuchians and goniopholidids and has never been described in those derived neosuchians considered close to eusuchians. A comparable depression was reported in a dyrosaurid specimen (e.g., Martin et al., 2019). The femur is otherwise more slender in its proportions than that of a dyrosaurid and does not provide sufficient characteristics to assign it to a family-level taxon.

Further comparison with the femur of *Terminonaris robusta* shows that the proximal head of the Salazac femur is nearly perpendicular to the shaft whereas its angle is more open in the former (Wu et al., 2001a). *Terminonaris robusta* also lacks the depression on the anterolateral surface of the proximal head. The Salazac specimen belongs to a much larger animal than the Carniol specimen. Whether the femoral depression represents an as yet unknown apomorphy of a putative basal form with close affinities to dyrosaurids is something to inspect with future discoveries of Mid-Cretaceous forms.

The total length of the Salazac crocodylomorph is estimated at about 3.5 m based on the regression equation established on femoral length in modern *Alligator mississippiensis* (Farlow et al., 2005). Given that the relative limb proportions are unknown in the Salazac specimen and
because of its phylogenetic distance compared to *Alligator* on which the regression was established, such dimensions can only be considered tentative. For example, Farlow et al. (2005) underlined that total length estimate in the extant *Gavialis gangeticus* was over-estimated because the species is a poor land-dweller with proportionally short limbs. Nevertheless, the sigmoid shaped femur of the Salazac specimen together with its typical “crocodilian” morphology suggests that the calculated TL is a relatively good estimate.

## 4. Discussion

The two newly reported specimens from the Aptian–Albian interval of southeastern France were collected from marine deposits. Given their depositional context, it is plausible that the marine environment corresponds to their living environment. Morphological evidence supporting the environmental origin of the newly reported specimens is unfortunately lacking. However, one feature can be noted and concerns the ornamentation of the external surface of the Carniol specimen: the cupular pattern is usually prominent in freshwater forms, although pholidosaurids possess such an ornamentation and are sometime recovered in marine deposits (Wu et al., 2001a; Hua et al., 2007). However, as discussed by Wu et al. (2001a), the habitat preferences of *Terminonaris robusta* are unknown and could represent a floating carcass. Conversely, fully marine forms such as Thalattosuchia or Dyrosauridae display smooth external bone surfaces on their skull roof. Obviously, relying on bone ornamentation alone is insufficient for inferring the habitat of extinct crocodylomorphs. In the present case, the freshwater-washout hypothesis is indirect and is regionally supported by the fact that Albian marine deposits of the Vocontian Basin have already yielded carcasses of terrestrial forms such as the theropod dinosaur *Genusaurus sisteronis* (Accarie et al., 1995) and terrestrial plant remains. In sum, interpreting the living environment of the newly reported remains is impossible with the limited morphological evidence at hand.

Crocodylomorph remains from the Aptian–Albian interval of Europe are extremely scarce in marine deposits. The dentary piece from the ?Aptian–Albian of the Isle of Wight (Young et al., 2014) represents an exception, and as with the specimens described in the present study from SE France, the question of its allochtony remains. It should also be noted that the Kimmeridge Clay Formation (KCF) regionally produces a wealth of vertebrate remains (Martill et al., 2006) and could represent a source of reworked specimens in younger strata. Therefore, it should not be excluded that the specimen reported by Young et al. (2014) was reworked from the KCF into Aptian–Albian deposits. Another notable example from this time interval concerns the nearly complete skeletons of *Pietraroiasuchus ormezzanoi*, a small hylaeochampsid eusuchian (Buscalioni et al., 2011). Those specimens, from the Albian Plattenkalk of the Apennines in Italy, were washed into a paralic depositional environment (Graziano et al., 2024) and the proximity of their living environment together with the nature of the deposits might explain their completeness. All other more or less contemporaneous European neosuchians have been reported from slightly older continental deposits of the late Barremian–early Aptian of Belgium and Spain with *Bernissartia fagesii* (Dollo, 1883; Buscalioni and Sanz, 1990). Elsewhere in the world, contemporaneous neosuchians are known exclusively from continental deposits with *Pachycheilosuchus trinquei* from the Albian of Texas (Rogers, 2003), *Wannchampsus kirpachi* and *Tarsomordeo winkleri* from the Aptian of Texas (Adams, 2014, 2019) and perhaps *Rugosuchus nonganensis* from the latest Early Cretaceous of China (Wu et al., 2001b). But none of these taxa closely resemble the squamosal described herein from the Carniol locality. The Carniol squamosal most resembles that of a pholidosaurid. This observation is congruent with the fossil record of the group that spans the latest Jurassic–early Late Cretaceous (Turonian) interval and includes members recovered both from freshwater and marine deposits (e.g., Wu et al., 2001a; Hua et al., 2007).

The observed temporal gap in the Aptian–Albian record of European crocodylomorphs frames several important faunal turnovers. For example,

in the marine environment, the last metriorhynchoids did not extend beyond the Valanginian (Sachs et al., 2020) and the next marine crocodylomorph occurrences are those of the eusuchian *Portugalosuchus azenhae* during the Cenomanian (Mateus et al., 2019) and of the pholidosaurid *Oceanosuchus boecensis* during the Cenomanian also (Hua et al., 2007). There are no confirmed dyrosaurids in the European fossil record but it should be noted that the group appears during the Campanian in Africa (Saber et al., 2025). In European freshwater deposits, the last goniopholidids are from the Albian of Spain (Buscalioni et al., 2013) and it is not until the Santonian–Campanian that eusuchians began to fill similar niches in freshwater ecosystems (Ösi et al., 2007; Martin and Buffetaut, 2008). Those latter forms have a phylogenetic tie to Barremian eusuchians such as *Hylaeochampsa vectiana* or *Bernissartia fagesii* and the Aptian–Albian record might be key to untangle relationships, as exemplified by *Pietraroiasuchus ormezzanoi* (Buscalioni et al., 2011). The phylogenetic relationships of atoposaurids with derived neosuchians are unsettled (see review in Pochat-Cottilloux et al., 2024), so any new Middle Cretaceous discoveries are potentially important in order to continue assessing phylogenetic relationships between Early Cretaceous (e.g., *Theriosuchus*) and Late Cretaceous (e.g., *Sabresuchus*) forms. Similarly, phylogenetic debates over the interrelationships of other neosuchian lineages may not be solved without access to forms from the Aptian–Albian interval (e.g., Tethysuchia vs Coelognathosuchia Young et al., 2014; Martin et al., 2014b; Kuzmin et al., 2025). Although the present report falls short of informative morphological characters, we recognise the potential of Aptian–Albian deposits to yield additional crocodylomorph material relevant to those questions.

### CRediT authorship contribution statement

**Jeremy E. Martin:** Writing – review & editing, Writing – original draft, Investigation, Conceptualization. **Romain Jattiot:** Writing – review & editing, Investigation.

### Declaration of competing interest

The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper.

### Acknowledgements

JEM thanks H. Capetta for raising awareness to the Carniol specimen and M. Mouana, A-L. Charruault and S. Adnet for help in accessing the specimen. JEM thanks Ryan McKellar (Royal Saskatchewan Museum) for providing pictures of the femur of *Terminonaris robusta*. The last version of the manuscript benefited from the constructive comments of two anonymous reviewers.

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