Scharwz-Wings et al, 2009

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Cretaceous Research 30 (2009) 1345–1355

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Cretaceous Research

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Lower Cretaceous Mesoeucrocodylians from Scandinavia(Denmark and Sweden)

Daniela Schwarz-Wings a,*, Jan Rees b, Johan Lindgren c

a Museum fur Naturkunde, Invalidenstrasse 43, D-10115 Berlin, Germanyb Soldattorpet 48, SE-653 50 Karlstad, Swedenc Department of Geology, GeoBiosphere Science Centre, Lund University, Solvegatan 12, SE-223 62 Lund, Sweden

a r t i c l e i n f o

Article history:Received 19 January 2009Accepted in revised form 22 July 2009Available online 30 July 2009

Keywords:ArchosauriaNeosuchiaBernissartiaGoniopholisPholidosaurusTheriosuchus

* Corresponding author.E-mail address: [email protected]

0195-6671/$ – see front matter � 2009 Elsevier Ltd.doi:10.1016/j.cretres.2009.07.011

a b s t r a c t

The crocodyliform faunas of the lowermost Cretaceous Rabekke and Jydegård Formations on the Balticisland of Bornholm, Denmark, and the Annero Formation of Skåne, southernmost Sweden, are repre-sented by isolated teeth, osteoderms, and vertebrae. The rich Berriasian assemblage of the RabekkeFormation includes at least three distinctive taxa: Bernissartia sp., Theriosuchus sp., and Goniopholis sp.,an association that is also known from several other contemporaneous European vertebrate localities. Incontrast to this fauna, the Jydegård and Annero Formations have yielded only rare mesoeucrocodylianremains, which are assigned to Theriosuchus sp. and an undetermined mesoeucrocodylian taxon, possiblyPholidosaurus. Geographically, the Scandinavian localities represent the easternmost and northernmostdistribution of typical continental Jurassic-Cretaceous crocodyliform communities in Europe.

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1. Introduction

Crocodyliform material from Scandinavia is generally scarce,with most finds comprising isolated, presumably shed teeth. Threenotable exceptions are the type and a referred specimen of Thor-acosaurus scanicus Troedsson, 1924 from the Danian (lower Paleo-cene) of southwestern Skåne (the southernmost province ofSweden) and an association of cranial elements from a singleindividual of Aigalosuchus villandensis Persson, 1959 from the lowerCampanian of the Kristianstad Basin, northeastern Skåne. Besidesrepresenting the most complete material of ancient crocodiliansfrom Scandinavia thus far, these specimens are the only ones thathave been described formally. Nonetheless, isolated teeth, verte-brae and osteoderms are occasionally found in Cretaceous strata ofsouthern Sweden (e.g., Siverson, 1993; Rees, 2002) and on theDanish island of Bornholm (Bonde, 2004; Lindgren et al., 2004,2008; Rees et al., 2005). Here, we present a description of some ofthese remains, focusing on finds from the Berriasian-Valanginianinterval of the Lower Cretaceous.

lin.de (D. Schwarz-Wings).

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2. Geological setting and localities

2.1. Bornholm, Denmark

On the Baltic island of Bornholm, lowermost Cretaceous sedi-ments are exposed in coastal cliff sections and a few inland quarrieson the southwestern part of the island (Fig. 1). The Cretaceous strataare included in the Nyker Group and the stratigraphic sequencespans the Berriasian-Valanginian interval (Gravesen et al., 1982).This group is subdivided into three consecutive units, denominatedfrom bottom to top the Rabekke, Robbedale, and Jydegård Forma-tions. The sedimentary sequence begins with fluvial and limnicdeposits of the Rabekke Formation, which are replaced upsectionby swamp and lake sediments of the same formation. These accu-mulations are in turn overlain by coastal and shallow marine sandsof the Robbedale Formation. Sediments at the top of the strati-graphic column are included in the Jydegård Formation, and consistprimarily of back-barrier and lagoonal sands (Noe-Nygaard et al.,1987; Noe-Nygaard and Surlyk, 1988).

2.1.1. Rabekke FormationThe outcrop producing crocodyliform remains is located on the

south coast of Bornholm, approximately one km east of the hamletof Arnager (Fig. 1). The sediments at this cliff section include mainlycoal-rich, black clays and silts interbedded with a few distinct lens-shaped beds of light grey to almost reddish clay stone. They were

Fig. 1. Map showing the location of Bornholm and Skåne in Scandinavia and detailed maps with site information for crocodyliform-yielding localities. 1, locality in the JydegårdFormation; 2, locality in the Rabekke Formation; 3, locality in the Annero Formation.

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probably deposited in a limnic environment with occasionalmarine influx, resulting in brackish water conditions (Petersenet al., 1996). Ostracode and palynomorph biostratigraphy indicatesthat the strata are early Berriasian in age, corresponding to theRunctoni ammonite Zone (Christensen, 1974; Lindgren et al., 2008,Fig. 2). A small number of lenses were sampled for micro-vertebrates, but only one yielded identifiable remains. In additionto teeth and bones of crocodyliforms (which are by far the mostcommon fossils at the site), the lens has produced teeth of sela-chians and actinopterygians, skeletal elements of amphibians andlizards (Rees et al., 2005), carapace fragments of turtles, teeth ofcarnivorous dinosaurs (Lindgren et al., 2008), and a single multi-tuberculate mammal tooth (Lindgren et al., 2004).

2.1.2. Jydegård FormationIn the lower part of the uppermost Berriasian-lower Valanginian

(Gravesen et al., 1982; Piasecki, 1984) Jydegård Formation (asexposed at A/S Carl Nielsen’s sand pit near Robbedale; Fig. 1) thereare two bivalve and gastropod mass mortality horizons (Noe-Nygaard et al., 1987; Noe-Nygaard and Surlyk, 1988). Sieving of therelatively coarse sands close to these horizons yielded a richmaterial of vertebrates, including numerous teeth of hybodontsharks and bony fishes, two teeth of dinosaurs (Bonde and Chris-tiansen, 2003; Christiansen and Bonde, 2003), a dentary of a smalllizard (Rees, 2000), and dentary fragments of crocodyliforms.

2.2. Skåne, Sweden

Lowermost Cretaceous deposits are normally not exposed inSkåne (due to a thick cover of Quaternary deposits), but in recentyears have been temporarily accessed through excavations near thehamlet of Eriksdal (Fig. 1). Due to tectonic activity in the VombTrough (Fig. 1), the Cretaceous strata in the area are tilted and

slightly overturned (e.g., Norling et al., 1993). Sediments accumu-lated during the latest Jurassic and earliest Cretaceous are includedin the Annero Formation, and were deposited in brackish waterenvironments with occasional fresh water influx (e.g., Guy-Ohlsonand Norling, 1994). The formation is subdivided in three consecu-tive members: Fyledal Clay (lower), Nytorp Sand (middle), andVitaback Clays (Erlstrom et al., 1991). The precise stratigraphicposition of the Jurassic-Cretaceous boundary is not known becauseage-diagnostic fossils are rare. However, it is assumed to be locatedwithin the lowermost part of the Vitaback Clays (Erlstrom et al.,1991), or, possibly, in the underlying Nytorp Sand (Vajda andWigforss-Lange, 2006). Rare crocodyliform teeth have beenrecovered from three beds belonging to the Vitaback Clays, i.e. VC3,VC7 and VC11 of Rees (2002). The lower bed (VC3) containsmolluscs indicative of mesohaline conditions, and the sedimentsmay originate from a lagoonal setting (Rees, 2002). Alternatively,this part of the Vitaback Clays represents a tsunami deposit (Vajdaand Wigforss-Lange, 2006). Higher up in the sequence, VC7 andVC11 were probably deposited in a small pond located further fromthe shoreline (Rees, 2002).

3. Material and methods

Sediments dominated by silts and clays (e.g., from Rabekke)were dried, soaked in a solution of sodium carbonate and hot water,and then washed through sieves with a mesh width of 0.500 mmand 0.355 mm, respectively. Sampled coquina beds (from Vitaback)were dissolved in buffered acetic acid, and magnetic as well asdensity separations were undertaken in order to concentrate thevertebrate remains (for more detailed information, see Rees, 2002).All residues were searched under a binocular microscope. Thecoarse sands of the Jydegård Formation were wet-sieved at thelocality using a net with a mesh width of 2 mm.

Fig. 2. Teeth of Bernissartia from the Berriasian Rabekke Formation of Bornholm, Denmark. A–C, MGUH 29103, 29104 and 29105, inflated, kidney-shaped teeth in labial, lingual,lateral, and apical views. D, MGUH 29106, kidney-shaped tooth in labial, lingual, and apical views. E, MGUH 29107, button-shaped tooth, enamel partly chipped away, in labial,lingual, lateral and apical views. F, MGUH 29108, button-shaped, heavily worn tooth, from left to right in labial, lingual, and apical views. All specimens are provided as SEMphotographs. Scale bars are 0.5 mm.

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3.1. Institutional abbreviations

BMNH; Natural History Museum, London (Former: BritischMuseum of Natural History).

MGUH; Natural History Museum, Copenhagen, Denmark.LO; Department of Geology, GeoBiosphere Science Centre, Lund

University, Lund, Sweden.

4. Systematic palaeontology

Crocodylomorpha Walker, 1970 (sensu Clark, 1986)Crocodyliformes Hay, 1930 (sensu Clark, 1986)Mesoeucrocodylia Whetstone and Whybrow, 1983 (sensuBenton and Clark, 1988)Neosuchia Benton and Clark, 1988Bernissartia Dollo, 1883

Bernissartia sp. indet.Fig. 2

Horizon and locality. Skyttegård Member, Rabekke Formation,Bornholm, Denmark.

Material. 10 teeth (MGUH 29103 - 29108, and 4 unnumberedspecimens, Natural History Museum, Copenhagen, Denmark).

Description. The height and width of the tooth crowns rangefrom 0.9 to 2.1 mm. Most of the crowns are inflated and distinctlylabio-lingually compressed (Fig. 2A–D). Near the transition to theroot, the base of the tooth crowns is constricted (Fig. 2A,B). Incross-section, the teeth are kidney-shaped, with a convex labialface and a slightly concave lingual surface (Fig. 2A–D). The labialand lingual faces are ornamented with apicobasally directed striaethat are evenly distributed over the apical two-thirds of thecrowns (Fig. 2B–D). Blunter teeth have more irregularly spacedstriations, occasionally shaped as inverted ‘Y’ (Fig. 2A). The apex ofunworn crowns is gently rounded (Fig. 2A,B), whereas abradedteeth possess a prominent wear-facet that is elliptical in occlusalview (Fig. 2C,D). With two exceptions (Fig. 2E,F), the teeth lackcutting edges (carinae), instead, the mesial and distal margins arebroadened.

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Remarks. The dentition of Bernissartia comprises conical teeth inthe rostralmost part of the snout and mandibles, broad and inflatedteeth in the mid-jaws, and button-shaped teeth in the caudal partof the dental arcade (Buffetaut, 1975; Buffetaut and Ford, 1979;Brinkmann, 1992). Enamel ornamentations are largely restricted tothe apical half of the tooth crowns, and range from vertical striae inkidney-shaped teeth to more or less unordered wrinkles in button-shaped ones.

The characteristic kidney-shaped tooth morphology of Bernis-sartia is not known from any other contemporaneous croco-dylomorph, and thus the teeth from Bornholm can be confidentlyassigned to that genus. The teeth of Bernissartia are distinguishedfrom those of Theriosuchus by being less labiolingually compressed,lacking distinct carinae, and possessing vertical or near verticallingual striae. Judging from the small size of the teeth from Rabekke,they may belong to juvenile individuals (Buscalioni and Sanz,1990b).

Atoposauridae Gervais, 1871Theriosuchus Owen, 1879

Theriosuchus sp. indet.Fig. 3

Horizons and localities. Skyttegård Member, Rabekke Formation,Bornholm, Denmark, and Vitaback Clays (Bed VC3 of Rees, 2002),Annero Formation, Skåne, Sweden.

Material. 284 teeth (LO 10485, MGUH 29109 - 29117, and 274unnumbered specimens, Natural History Museum, Copenhagen,Denmark).

Description. Two crown morphotypes are here assigned to Ther-iosuchus. The first morphotype includes 184 teeth with a lanceolateand labiolingually compressed crown shape (Fig. 3A–D). The teethvary considerably in size; from 1 to 2 mm in basal crown length andfrom 1 to 4 mm in total crown height. The tooth crowns are con-stricted at their base, and the base itself is slightly inflated (Fig. 3B–D). Moreover, the tooth crowns bear mesial and distal carinae. Thelabial face of the teeth is strongly convex, and the apical half of thissurface exhibits vertical striae (Fig. 3A–D). The lingual side of thecrowns is weakly convex, and occasionally possesses a shallowcentral groove on its basal third (Fig. 3C). The enamel on this side isornamented with more pronounced striae than on the labial face.The striations may be either closely (Fig. 3A,B) or widely (Fig. 3C,D)spaced. Whereas the striae in the central two-thirds of the toothcrowns extend more or less apicobasally, they fan out in the mesialand distal parts of the crowns to terminate at or near the carina,giving the latter a roughened appearance.

The second morphotype comprises 99 teeth with mesiodistallybroad crowns and a weakly convex to almost horizontal apicalmargin (Fig. 3E–H). There is a great size variation within this group,with both lengths and heights ranging from 1 to 3 mm. All teeth arestrongly labiolingually compressed with a weak constriction nearthe base of the crowns. Carinae are present along the mesial anddistal margins. In its lower half, the labial surface of the crowns isstrongly convex, whereas the lingual surface is less so. The enamelis developed into regularly distributed striae on both the labial andlingual surfaces. The distribution of striae is similar to that of thelanceolate morphotype; on the labial side, the striae extend regu-larly from the base to the apex of the crowns, whereas on thelingual side, centrally located striae converge distally and terminatenear the apex. Marginal striations on the lingual face are directedtoward the carinae, resulting in a jagged appearance (Fig. 3E,H).

Within the referred material there is also one small-sized toothMGUH 29117 (0.8 mm in maximum length and less than 1 mm inoverall height), presumably from an immature individual, thatpreserves large parts of the root (Fig. 3I). Although incomplete, the

preserved parts suggest that the root was originally barrel-shaped.The tooth crown is inflated at its base and labiolingually flattened.The crown is rounded in labial and lingual aspects and possessesmesial and distal carinae. The enamel forms a small number ofwidely spaced striae, which are more or less apicobasally directed.

Remarks. The dentition of Theriosuchus includes three differentmorphotypes (Owen, 1879; Joffe, 1967; Brinkmann, 1992; Salisbury,2002; Schwarz and Salisbury, 2005): (i) slender and conical teethwhere the striations are largely restricted to the lingual face of thecrowns, found in premaxillary, rostral-most maxillary, and rostral-most dentary positions (Owen, 1879; Brinkmann, 1992; Schwarzand Salisbury, 2005); (ii) lanceolate teeth with a fan-shapeddistribution of the marginal lingual striae are found in the middleand caudal portions of the jaws (Brinkmann, 1992; Owen, 1879;Schwarz and Salisbury, 2005); and (iii) a third morphology is foundin Theriosuchus pusillus and T. ibericus, where the teeth are broadand strongly labiolingually compressed, and both the lingual andlabial faces are covered with striations (although fan-shaped striaeare present only on the lingual face) (Owen, 1879; Brinkmann,1989; Brinkmann, 1992; Salisbury, 2002). Generally, themorphology of the teeth of Theriosuchus varies along the dentalarcade, with the lanceolate type becoming proportionately broaderand slightly more rounded towards the caudal end of the jaws.Dentary teeth possess stronger striations than do maxillary teeth.In addition, there may also be some ontogenetic variation, becausethe striations are more strongly developed in older (larger) indi-viduals than they are in younger (smaller) ones.

The teeth from Bornholm and Skåne are assigned to Ther-iosuchus on the basis of their overall shape and the configuration ofthe striae. Whereas a lanceolate shape is commonly occurring inTheriosuchus (Owen, 1879; Joffe, 1967; Buffetaut, 1983; Brinkmann,1989, 1992; Thies et al., 1997; Winkler, 1995; Salisbury, 2002;Schwarz and Salisbury, 2005), broad and labiolingually compressedcrowns have previously been found only in T. pusillus and T. ibericus.The morphology of MGUH 29117 slightly derives from the typicaltooth morphology of Theriosuchus, which may be a result ofontogeny, or it may also represent another species within theAtoposauridae, such as Montsecosuchus (Buscalioni and Sanz,1990a).

Goniopholididae Cope, 1875Goniopholis Owen, 1842

Goniopholis sp. indet.Fig. 4A–C

Horizon and locality. Skyttegård Member, Rabekke Formation,Bornholm, Denmark.

Material. Five teeth (MGUH 29118 - 29120 and two unnumberedteeth, Natural History Museum, Copenhagen, Denmark).

Description. Five conical tooth crowns from Rabekke are hereassigned to Goniopholis. The teeth range from 1 to 4 mm in basalcrown length and from 2 to 8 mm in total crown height. They are onlymodestly compressed labiolingually, resulting in an almost circularbase (4A–C). The crowns are slightly recurved, taper apically, and endin a rounded apex. With increased size, the teeth become relativelybroad and thickset; smaller teeth have a width-to-height ratio of 1:3,whereas larger teeth have a corresponding ratio of 1:2. Unserratedmesial and distal carinae are present, although they are poorlydeveloped. The enamel forms rounded ridges that extend verticallyover the entire height of the crowns, although they weaken some-what towards the apex (Fig. 4A,C). In other teeth, the ridges arerestricted to the centre of the crowns (Fig. 4B).

Remarks. Teeth of Goniopholis are stout and conical, and there islittle morphological change along the jaw (e.g., Hulke, 1878; Owen,

Fig. 3. Teeth of Theriosuchus from the Berriasian Rabekke Formation of Bornholm, Denmark. A, MGUH 29109, conical, nearly lanceolate tooth, in labial, lingual, lateral, and apicalviews. B, MGUH 29110, lanceolate tooth, in labial, lingual, and lateral views. C–D, MGUH 29111 and 29112, lanceolate teeth, in labial, lingual, lateral, and apical views. E–F, MGUH29113 and 29114, broadened teeth, in labial, lingual, lateral, and apical views. G, MGUH 29115, broadened tooth, in labial, lingual, and apical views. H, MGUH 29116, broadened tooth,in labial, lingual, and lateral views. I, MGUH 29117, tooth of a presumably early juvenile individual, in labial, and lingual views. All specimens are provided as SEM photographs. Scalebars for A–H are 0.5 mm, scale bar for I is 0.1 mm.

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1878, 1879; Dollo, 1883; Koken, 1887; Hooley, 1907; Ortega et al.,1996; Krebs and Schwarz, 2000; Salisbury, 2002; Schwarz, 2002).The teeth are slightly labiolingually compressed, and occasionallybear weak carinae. In smaller tooth crowns, the enamel striationsare coarse and well-spaced from one another, whereas they mayoverlap, and become weaker and/or more closely spaced on largerteeth. A small and presumably juvenile specimen referable toGoniopholis gracilidens (Salisbury, 2002) (BMNH No. 48217, ‘Nan-nosuchus gracilidens’ as characterized by Owen, 1879) possessesconical but slender and strongly curved teeth (Owen, 1879; Joffe,1967), suggesting that there may be ontogenetic differences withinGoniopholis. However, the specimen is considered to represent

a species different from all other described Goniopholis species fromthe Purbeck Formation (Salisbury, 2002).

The stout proportions and coarse striations on the teeth fromBornholm suggest that they belong to Goniopholis (e.g., Owen, 1878,1879; Krebs and Schwarz, 2000; Salisbury, 2002). The teeth aredistinguishable from the more slender and finely striated teeth ofPholidosaurus, which has previously been reported from slightlyyounger strata on this island (Bonde, 2004), and they differ alsofrom the needle-like and strongly recurved teeth of ‘Nannosuchus’.

Mesoeucrocodylia indet.Figs. 4E–I, 5A–F

Fig. 4. Teeth of Goniopholis from the Berriasian Rabekke Formation of Bornholm, Denmark. A, MGUH 29118, conical tooth in labial, lingual, apical, and lateral views. B, MGUH 29119,conical tooth in labial, lingual, and apical views. C, MGUH 29120, conical tooth in labial, and lingual views. D, MGUH 29127, undetermined isolated tooth of an undeterminedarchosaur from Bornholm, in labial, lingual, lateral, and apical views. E, MGUH 29127, undetermined conical tooth of mesoeucrocodylian from Bornholm, in labial, lingual, apical, andlateral views. Postcranial remains of mesoeucrocodylians from Bornholm. F–G, MGUH 29122 and 29123, centra of dorsal vertebrae in left lateral (above) and caudal (below) view. H,MGUH 29125, fragmentary osteoderm in external (left) and internal (right) views. I, MGUH 29124, incomplete vertebral centrum in cranial view. Specimens in A-E are provided asSEM photographs, specimens in F–I have been produced with a multi-focus camera. Scale bar for A, F, G, and I is 1 mm, scale bars for B-E are 0.5 mm, scale bar for H is 2 mm.

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Horizon and locality. Skyttegård Member, Rabekke Formation,Bornholm, Denmark.

Material. 180 teeth (MGUH 29121 and 179 unnumber speci-mens), three vertebral centra (MGUH 29122 - 29124), and 24fragmentary osteoderms (MGUH 29125, and 23 unnumberedspecimens).

Description. TeethdThe tooth crowns are slender, conical andsomewhat compressed labiolingually. The apex is pointed andslightly curved lingually. Weak carinae extend from the base to theapex of the crowns. The enamel is ornamented with vertical striaethat increase in size and number with increased tooth size (Fig. 4E).

VertebraedThe assemblage from the Rabekke Formationincludes two isolated yet virtually complete vertebral centrameasuring about 4.5 mm in length (Fig. 4F,G), in addition toa partial vertebra of similar size (Fig. 4I). The parapophyses of one ofthe vertebrae (MGUH 29122) are positioned craniodorsally andcover the neurocentral suture (Fig. 4F), indicating that this isa cranial thoracic vertebra. The other vertebral centrum (MGUH29123) does not possess parapophyses, and thus probably origi-nates from the mid-thoracic region (Fig. 4G). Both vertebrae areamphicoelous with a well-developed concavity at both the cranialand caudal articular surface. The vertebral centra are hour-glass-

Fig. 5. Teeth and osteoderm of mesoeucrocodylians from the BerriasianVitaback Clays of Skåne, Sweden. A-B, undetermined isolated teeth (LO 10485) from VC3, C, Undeterminedisolated tooth (LO 10486) from VC7, D, Undetermined isolated tooth (LO 10487) from VC11. Isolated incomplete osteoderm with partial dorsal keel (LO 10488) from VC11 in E,external and F, internal view. All pictures produced with a multi-focus camera. Scale bar for A–B and D–F is 2 mm, scale bar for C is 1 mm.

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shaped, and their lateral surfaces are perforated by nutritiveforamina.

OsteodermsdAlthough incomplete, the osteoderms probablyhad a sub-rectangular outline. The external surface is sculpturedwith circular pits (Fig. 4H), and an incomplete median keel ispreserved in a few specimens. The internal surface is smooth(Fig. 4H).

Remarks. Conical and striated teeth are present in the rostralportions of the jaws in Theriosuchus, Bernissartia (Dollo, 1883;Buffetaut and Ford, 1979; Buscalioni et al., 1984), and a juvenilespecimen of Goniopholis (‘Nannosuchus’) gracilidens (Owen, 1879;Joffe, 1967; Salisbury, 2002). These teeth are known to vary inmorphology during different ontogenetic stages, and are thusinsufficient for generic determination. Although the teeth underdiscussion most likely originate from one or more of the abovementioned taxa, it is currently not possible to exclude affinitieswith other contemporary mesoeucrocodylians with slender andconical teeth, such as various thalattosuchians and pholidosaurids(Buffetaut, 1982; Hua 1997; Hua and Buffetaut, 1997; Bonde, 2004;Pouech et al., 2006).

The amphicoelous vertebral centra correspond morphologicallyto thoracic vertebrae of Theriosuchus pusillus (Owen, 1879; Salisburyand Frey, 2001) and T. guimarotae (Schwarz and Salisbury, 2005).However, because postcrania of juvenile Goniopholis are unknown,and because the vertebrae of adult individuals possess no taxo-nomically useful characters (Owen, 1878; Salisbury, 2001; Schwarz,2002), it is not possible to exclude an affinity with e.g., Goniopholis.Similarly, the thoracic vertebrae of Bernissartia (Buscalioni and Sanz,1990b; Norell and Clark, 1990; Salisbury and Frey, 2001) are notsignificantly different from those of Theriosuchus.

Osteoderms may be distinguished by their shape, size, presenceor absence of one or two external keels, and shape of the cranialarticular peg. However, because none of these characters arepreserved in the osteoderms from Rabekke, an unambiguoustaxonomic assignment is currently not possible.

Horizon and locality. Tornhøj Member, Jydegård Formation,Bornholm, Denmark.

Material. Nine mandibular fragments (MGUH 29126), presum-ably from a single individual.

Description. The fragments are all of comparable size and likelyto originate from a single mandible, although they do not fittogether. The most complete specimen (preserved length is 45 mm)is a post-symphyseal dentary fragment with two alveoli. Aspreserved, it is 19 mm high, although the bone is incompleteventrally. The dorsal and lateral margins are straight. Two alveoli

are widely separated from one another and are sub-circular inoutline. The broken fragment of a tooth is visible in one of thealveoli, exposing a circular tooth base with a medially open pulpcavity. The lateral surface of the bone is sculptured with shallow,longitudinally directed grooves, whereas the medial face is brokenoff, exposing parts of the Meckelian canal and the bases of thedentary alveoli.

Remarks. There are no taxonomically useful characterspreserved in the mandibular material, rendering it impossible todetermine whether it belongs to a large individual of one of thecrocodyliforms described above, or represent another, larger taxon.

Horizon and locality. Vitaback Clays (Bed VC3, VC7 and VC11 ofRees, 2002), Annero Formation, Skåne, Sweden.

Material. Four teeth (LO 10485–10487) and two osteoderms (LO10488, and one unnumbered specimen, Department of Geology,Lund University).

Description. TeethdThree conical, pseudocaniniform teeth havebeen found in VC3. Two of the teeth include parts of the root(Fig. 5A,B), whereas the third, and much smaller specimen, onlycomprises the crown. The smallest tooth crown has a basal diameterof 1 mm and a height of 2 mm, while the two larger teeth measure3 mm in basal diameter and 8 mm in height. The tooth crowns arecurved medially and lack basal constrictions. They are somewhatcompressed mesiodistally. The teeth taper apically and end ina narrow yet rounded apex, which is clearly offset from the rest of thecrown. Carinae are weakly developed on the distal (?) margin of thecrowns. The enamel forms distinct ridges, which are widely spacedand extend apicobasally. At the base of the tooth, the ending of theglossy enamel marks the transition to the root. A tooth crown witha similar morphology was found in VC11 (Fig. 5D), in addition toa very small (2 mm high) conical tooth that lacks striations.

An incomplete conical tooth crown (apex missing) with a heightof 5 mm was found in VC7. The crown is stout and bears mesial anddistal carinae (Fig. 5C). With the exception of a few wrinkles on theapical half of the crown, the enamel is virtually smooth.

OsteodermsdTwo incomplete and externally sculptured dorsalosteoderms are morphologically similar to the correspondingelements from the Rabekke Formation. One of the specimensappears to have had an oval outline, indicating that it is a caudal, orpossibly a nuchal osteoderm. The other osteoderm has a sub-rect-angular shape and possesses remnants of a medial keel, suggestingthat it is part of a dorsal or caudal osteoderm (Fig. 5E,F). The internalsurface is smooth in both specimens.

Remarks. The isolated conical teeth from the Vitaback Clays can,because of their stoutness, the presence of coarse striae, distal

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carinae and a distinct smooth tip, be excluded from Pterosauria. Thepresence of carinae and the widely spaced striations excludes themalso from Plesiosauria. The isolated teeth resemble a tooth tenta-tively assigned to Pholidosauridae by Bonde (2004). However, thereare some differences with respect to the coarseness and amount ofthe striae, which prevent a confident assignment.

Archosauriformes Gauthier, 1986Archosauria Cope, 1869

Archosauria indet.Fig. 4D

Horizon and locality. Skyttegård Member, Rabekke Formation,Bornholm, Denmark.

Material. One tooth (MGUH 29127).Description. The tooth crown has a length of 2.2 mm and a height

of 1.6 mm. It is strongly compressed labiolingually and has a leaf-shaped lateral outline with convex mesial and distal margins(Fig. 4D). The carinae are limited to the apical two-thirds of thecrown and bear distinct denticles. In occlusal view (Fig. 4, right handillustration), the cutting edge is flexed labially and lingually into an‘S’-shaped structure. The apex of the crown is formed by an enlargeddenticle. The base of the tooth crown is inflated and less constrictedthan in the teeth described above. The enamel is smooth.

Remarks. MGUH 29127 cannot be unambiguously assigned toany known mesoeucrocodylian. Denticulated carinae occur also ina number of archosaurian groups including ziphosuchian croco-dyliforms, some Triassic pterosaurs (Dalla Veccia, 2003) and dino-saurs (e.g., Farlow et al., 1991; Holtz et al., 1998; Ortega et al., 2000).In contrast, some other vertebrate groups, such as varanids andmosasaurid squamates and fishes, possess carinae with fine serra-tions, but no distinct denticles are formed at the cutting edges(compare e.g., Jagt et al., 2005). Triassic basal archosaurs such asEuparkeria (Ewer, 1965), erythrosuchids (e.g., Modesto and Botha-Brink, 2008), phytosaurs (e.g., Hungerbuhler, 2000), or raui-suchians (e.g., Nesbitt, 2005) can be ruled out because of theirconsiderably older appearance and different, blade-like toothmorphology. Theropod dinosaurs can equally be ruled out by theirsignificant differences in tooth morphology (Farlow et al., 1991;Abler, 1992; Holtz et al., 1998), which is also the case for the leaf-shaped, denticulate teeth of prosauropods (e.g., Barrett, 2000). Theoverall crown morphology and regular denticles are consistentwith teeth of the ornithischian (?) dinosaur Pekinosaurus (Hunt andLucas, 1994). However, this taxon is reported from the UpperTriassic of North Carolina (USA), and is therefore considerably olderthan the specimen from Bornholm. Among non-eusuchian meso-eucrocodylians, denticulated serrated teeth occur in Theriosuchusibericus (Brinkmann 1989, 1992), Dakosaurus andiniensis (Gaspariniet al., 2006), and in the Ziphosuchia (Ortega et al., 2000). Theoverall shape and serration profile of MGUH 29127 correspondsroughly with those of T. ibericus and the ziphosuchian ‘‘Araripe-suchus’’ wegeneri (Ortega et al., 2000) from the Lower Cretaceous ofNiger, although it lacks enamel striations. It is thus possible thatMGUH 29127 represents a taxon distinct from the other describedcrocodyliforms, another species of Theriosuchus, or an unidentifiedziphosuchian.

5. Taphonomy and size estimates

The majority of the crocodyliform teeth from the RabekkeFormation and Vitaback Clays are well preserved and show little orno sign of transportation. However, they lack roots and have oftenworn and broken apices, indicating that they were lost during thenormal process of tooth replacement (Kieser et al., 1993). The

presence of replacement teeth indicates that the localities weresituated close to the habitat of these crocodyliforms. A few teethshow some abrasion, suggesting that they were transported oversome distance. Transportation may also have damaged the verte-brae by separating, at least in immature individuals, the neuralarches from the centra. However, the method of collecting thesefossils by sieving and chemical preparation might also havecontributed to abrasion and breakage. It is thus possible that partsof the material at hand were embedded after a short transportationfrom the surrounding environment, and other parts wereembedded within the environment.

The crocodyliform teeth and bones from the Rabekke Formationare generally small and cannot have belonged to animals largerthan about 60 cm in total body length. This is in accordance withthe estimated size range of Theriosuchus, which reached a length ofabout 50 cm (Schwarz and Salisbury, 2005), and Bernissartia, witha body length of some 60 cm (Salisbury, 2001). The recoveredmaterial suggests that both taxa are probably represented by bothjuvenile and adult individuals. In contrast, Goniopholis could attainbody lengths in excess of three metres (Salisbury, 2001), andconsequently this taxon must be represented by juveniles only. Theabsence of adult specimens of Goniopholis may be a result ofenvironmental specificity linked to dietary changes from juvenileto adult animals (Cott, 1961; Trutnau, 1994), or could be a result oftaphonomic factors.

6. Palaeobiogeography

The crocodyliform assemblages from Bornholm and Skåne aresimilar in their composition of more than two fossil crocodyliformtaxa (i.e., Bernissartia, Theriosuchus, Goniopholis and additionally,small-sized taxa or a larger taxon such as Pholidosaurus) toa number of contemporaneous faunas of west-central Europe(Fig. 6), including:

(1) Guimarota in Leiria, Portugal; ‘‘Guimarota-Strata’’ of theAlcobaça-Formation, Kimmeridgian (Upper Jurassic). Partialskeletons and isolated dental and skeletal elements ofMachimosaurus hugii (Krebs, 1967, 1968), Bernissartia sp.(Brinkmann, 1989), Lisboasaurus estesi (Buscalioni et al., 1996),Goniopholis baryglyphaeus (Schwarz, 2002), Lusitanisuchusmitracostatus (Schwarz and Fechner, 2004), and Theriosuchusguimarotae (Schwarz and Salisbury, 2005).

(2) Langenberg/Oker in northwestern Germany; LangenbergFormation (Kimmeridgian). Partial skeletons of Theriosuchus,Goniopholis, Machimosaurus, and Steneosaurus (Thies et al.,1997; Thies and Broschinski, 2001; Karl et al., 2006).

(3) Andres in Pombal, Portugal; Alcobaça-Formation (Kimme-ridgian–Tithonian). Isolated skeletal remains of Theriosuchussp. and Goniopholis sp. (Malafaia et al., 2006).

(4) Boulogne-sur-Mer and Wimille, Boulonnais, France; Forma-tions greseuses, ‘‘Montrouge’’ and ‘‘LaRochette II’’ (Tithonian/Portlandian). Teeth and incomplete bones of Goniopholis cf.simus (Sauvage, 1874, 1882; Buffetaut, 1986; Salisbury et al.,1999) and Theriosuchus sp. (Cuny et al., 1991).

(5) Swanage, Dorset, Great Britain (several localities); PurbeckLimestone Group, ‘Beckles residuary marls’ (Salisbury, 2002)(Berriasian). Partial skeletons and isolated teeth and bones ofTheriosuchus pusillus, Goniopholis crassidens, G. simus, Pholi-dosaurus purbeckensis (Salisbury, 2002), cf. Bernissartia(Owen, 1878, 1879; Salisbury, 2002), and cf. Lisboasaurus(Evans, 1994).

(6) Cherves-de-Cognac, France; Gypsum and marlstone unitsequivalent to Purbeck Limestone Group (Berriasian). Skeletonsof Theriosuchus spp., Goniopholis simus, G. crassidens, a single

Fig. 6. Geographic map with Upper Jurassic and Lower Cretaceous localities yielding mesoeucrocodylian communities with Goniopholis, Theriosuchus, and Bernissartia. Numbersrefer to section 6, Palaeobiogeography; crosses refer to the localities on Bornholm and Skåne described in this paper.

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skull of Pholidosaurus purbeckensis, isolated mandible of Ber-nissartia fagesi, isolated teeth of these taxa (Pouech et al.,2006; Mazin and Pouech, 2008; Mazin et al., 2006, 2008).

(7) Isle of Wight, Great Britain; Wessex Formation, Wealden(Hauterivian-Barremian). Cranial and postcrania ofGoniopholis crassidens (Owen, 1842; Hooley, 1907), teeth ofPholidosaurus sp. (Owen, 1842), teeth of Bernissartia sp.(Buffetaut and Ford, 1979), and a single skull of Theriosuchussp. (Buffetaut, 1983).

(8) Galve/Teruel, Spain; El Castellar Formation and CamarillasFormation (Hauterivian-Barremian). Skull and teeth ofBernissartia sp. (Buscalioni et al., 1984), teeth and bones ofTheriosuchus sp. (Buscalioni and Sanz, 1987b), and skeletalelements of Goniopholis (Buscalioni and Sanz, 1987a,b;Sanchez-Hernandez et al., 2007).

(9) Una/Cuenca, Spain; Una-Formation (Barremian). Partial skel-etons of Unasuchus reginae and Theriosuchus ibericus, isolatedskeletal elements of Goniopholis sp. and Bernissartia sp.(Brinkmann, 1989, 1992), dentary of Lisboasaurus (Schwarzand Fechner, 2008).

(10) Pio Pajaron,Cuenca, Spain; Una-Formation (Barremian). Teethof Theriosuchus ibericus and Bernissartia sp. (Winkler, 1995).

(11) Buenache de la Sierra, Spain; La Huerguina LimestoneFormation (Upper Barremian). Isolated teeth of cf. Goniopholisgracilidens, Theriosuchus sp., Bernissartidae indet., and Croc-odylomorpha indet. (Buscalioni et al., 2008).

(12) Vallipon and La Cantalera, Teruel, Spain; Artoles Member (Bar-remian-Aptian). Isolated remains of Goniopholis sp., Bernissartiasp., and Theriosuchus sp. (Ruiz-Omenaca and Canudo, 2001).

(13) Bernissart, Belgium; Sainte-Barbe clays, Wealden (Barremian-Aptian). Complete skeletons of Goniopholis sp. and Bernissartiafagesii (Dollo, 1883; Buffetaut, 1975; Salisbury et al., 1999).

Geographically, the Scandinavian localities represent the east-ernmost and northernmost distribution of typical continentalJurassic-Cretaceous crocodyliform communities in Europe (Fig. 6).During the Late Jurassic and Early Cretaceous, most parts of Europewere covered by shallow epicontinental seas (Ziegler, 1990; Smithet al., 1994). Continental islands existed, such as the Iberian Meseta,and the localities listed above were placed along their margins.Whereas the general south-to-north distribution and position ofthese localities (Fig. 6) was not different during the Cretaceous thankit is today, the whole area was situated farther to the south,stretching between a latitude of 30� at the southernmost Spanishlocality Buenache de la Sierra and of 45� at the northernmost localityin Skåne (Smith et al., 1994, and: http://jan.ucc.nau.edu/~rcb7/nat.html). The common crocodyliform genera Goniopholis and Ther-iosuchus have been documented partly by bone material, but also byfrequent findings of isolated teeth, and their occurrence in an areaslightly more north- and eastward than hitherto known is notsurprising. In contrast, Bernissartia is a rarer element, and its occur-rence on Bornholm extends the geographical range of this taxonconsiderably. The Berriasian age of the Bornholm and Skåne locali-ties demonstrates that already in the earliest Cretaceous, this kind ofcrocodyliform assemblage was far distributed, corresponding to thecoastal margins of the islands (Smith et al., 1994). Additionally tothese more common taxa, some of the localities listed above haveyielded evidence of very rare, or even endemic taxa: Lisboasaurus,

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Lusitanisuchus, and Unasuchus as well as a mixture of phylogeneti-cally divergent taxa (Buscalioni and Vullo, 2008). All of these local-ities are contrasted by the Late Jurassic ‘‘Plattenkalk’’ localities Cerin/France (Lortet, 1892) and Solnhofen in Germany (Wellnhofer, 1971),and the Berriasian ‘‘Plattenkalk’’ locality El Montsec in Spain (Bus-calioni and Sanz, 1990a), which yield crocodyliform assemblageswith exclusively small atoposaurid taxa, and in the case of Cerin, therelatively small mesoeucrocodylian Crocodileimus. A shift in thecrocodyliform faunas at the end of the Early Cretaceous causedreplacement of these faunal assemblages of non-neosuchian meso-eucrocodylians by eusuchian-ziphosuchian faunas (Buscalioni andVullo, 2008).

Acknowledgements

The fieldwork was carried out on a number of occasions withfinancial support from Lund Geological Field Club, the RoyalSwedish Academy of Sciences, the Natural History Museum inCopenhagen, and Nordisk Forskerutdanningsakademi (NorFa). Wethank Mikael Siverson and Regitze Benthien for assistance in thefield and for donating important specimens. DSW thanks GabrieleDrescher for her assistance with the SEM photographs, HenrikStohr for his assistance with the multi-focus camera, and OliverWings for discussion of the palaeogeographic situation. JR and JLacknowledge the Swedish Research Council (Vetenskapsrådet) forfunding their research. We thank Steven Salisbury and an anony-mous Cretaceous Research reviewer for their helpful comments.

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