• Campanian;
  • Ceratopsia;
  • Leptoceratopsidae;
  • Neoceratopsia;
  • Sweden;
  • teeth;
  • vertebrae


  1. Top of page
  2. Abstract
  3. Geological setting and localities
  4. Material and methods
  5. Systematic palaeontology
  6. References

Abstract:  Shallow marine, nearshore strata of earliest Campanian (Gonioteuthis granulataquadrata belemnite Zone) and latest Early Campanian (informal Belemnellocamax mammillatus belemnite zone) age in the Kristianstad Basin, southern Sweden, have yielded isolated leptoceratopsid teeth and vertebrae, representing the first record of horned dinosaurs from Europe. The new leptoceratopsid occurrence may support a European dispersal route for the Leptoceratopsidae, or may represent an entirely endemic population. The presence of leptoceratopsid teeth in shallow marine deposits contradicts previous hypotheses suggesting that basal neoceratopsians mainly preferred arid and/or semi-arid habitats far from coastal areas.

Ever since the discovery of isolated dental and skeletal remains in southern England almost 200 years ago (McGowan 2001), dinosaurs have attracted widespread attention, being some of the largest and most conspicuous land animals that have ever lived. Ceratopsia, or horned dinosaurs, represent one of the last major radiations of non-avian Dinosauria prior to the end of the Cretaceous. Systematically, this diverse assemblage of frilled herbivorous ornithischians is subdivided into the Psittacosauridae and Neoceratopsia, which, in turn, comprises the Ceratopsoidea (Ceratopsidae + Leptoceratopsidae), Protoceratopsidae and a number of primitive taxa, such as ArchaeoceratopsDong and Azuma, 1997 and LiaoceratopsXu et al., 2002 (Makovicky 2001; You and Dodson 2003). Whereas the monophyletic status of the Neoceratopsia and Ceratopsidae has been firmly established in a number of recent cladistic analyses, the Leptoceratopsidae and Protoceratopsidae are supported by only a few characters (Sereno 1986, 2000; Xu et al. 2002; You and Dodson 2003; Chinnery 2004).

Remains of horned dinosaurs have hitherto mainly been restricted to central Asia and western North America (Ryan and Currie 1998; but see also Chinnery et al. 1998), although fragmentary skeletal elements from the Aptian, Lower Cretaceous of Victoria, Australia, indicate that ceratopsians may also have been present in the Southern Hemisphere (Rich and Vickers-Rich 1994). Here, we report upon the first occurrence of neoceratopsians from Europe. The material comprises isolated teeth and vertebrae collected from shallow, nearshore marine strata of earliest Campanian and latest Early Campanian age (in belemnite terms; see Christensen 1975) in the Kristianstad Basin, southern Sweden. Despite the fragmentary nature of the material at hand, the neoceratopsian record from this basin has interesting biogeographical implications, and adds significantly to our knowledge of the function of the specialized mastication apparatus in the Leptoceratopsidae.

Geological setting and localities

  1. Top of page
  2. Abstract
  3. Geological setting and localities
  4. Material and methods
  5. Systematic palaeontology
  6. References

In the Kristianstad Basin, a deeply weathered, Precambrian crystalline basement with markedly uneven small-scale topography (Bergström and Sundquist 1978; Lidmar-Bergström 1982) is overlain by up to 250 m (Erlström and Gabrielson 1992) of predominantly shallow water, fully marine deposits of Barremian (Norling and Skoglund 1977; Bergström and Sundquist 1978; Norling 1981) to earliest Maastrichtian age (Siverson 1993b). At several localities in the north-eastern part of the basin, however, marine strata of latest Early Campanian age rest upon delta plain deposits of late Santonian/earliest Campanian age.

The crystalline basement surface slopes to the south and is cut off by the Linderödsåsen and Nävlingeåsen horsts, which together mark the south-western boundary of the basin. The south-eastern boundary is arbitrarily drawn along the shore of the Baltic Sea (see Kumpas 1980 for a different definition and nomenclature of the basin). The northern demarcation of the basin is rather diffuse with several small outliers of Upper Cretaceous (typically mid-Campanian or lowermost Maastrichtian) rocks.

Accessible strata exposed in numerous quarries and a few natural outcrops along the margins of the basin range in age from the Early or early Middle Santonian (Christensen 1975) to earliest Maastrichtian (Siverson 1993a; but see also Lindgren and Siverson 2002, p. 75). Tertiary strata are not known to be present in the Kristianstad Basin area, although loose blocks of Paleocene age have been found at the southern tip of the basin (Sandegren 1914).

Common, storm-generated conglomerates (with crystalline pebbles, belemnite guards and oysters) in the Lower Campanian part of the sequence attest to nearshore, high-energy environments (Erlström and Gabrielson 1986, 1992). These conglomerates are part of a broad spectrum of shallow water marine lithologies reflecting a mosaic of local environments within an archipelago setting. Calcareous, glauconitic quartz sands, calcarenites and oyster banks are the dominant sediment types in the marine Lower Campanian.


Åsen (marine sand unit; Text-fig. 1). Map sheet: Karlshamn 3E NV (topographical map sheet, 1 : 50,000); coordinates UTM VC 690 229 (56°08′56·1″N, 14°29′56·0″E). All four leptoceratopsid teeth (as well as the vertebrae) from Åsen are from the lower half of the section, which is latest Early Campanian in age (Christensen 1975; Siverson 1992; Rees 1999; Lindgren and Siverson 2002). The small, presumably juvenile maxillary tooth (RM PZ R1828; Text-fig. 3C–E) was found in a highly calcareous quartz sand with extremely abundant and well-preserved belemnite guards and articulated oysters, which makes up the uppermost 0·5 m of the uppermost Lower Campanian part of the sequence. The other three teeth (RM PZ R1827, 1829, 1830; Text-figs 2B, 3A–B, J–K) are all from a mottled, 10-cm-thick storm deposit comprising greenish, coarse quartz sand mixed with fragmented oysters, belemnites and common vertebrate remains. The bed is situated approximately 150 cm below the discontinuity surface in undisturbed (or composite) sections and rests upon coarse quartz sand in a grey fine matrix of reworked kaolin clay. It is overlain by green, fine, glauconitic quartz sand that is progressively more calcareous up-section.


Figure TEXT-FIG. 1..   Simplified geological map of southern Sweden showing the location of the Kristianstad Basin in the north-eastern corner of the province of Skåne and the localities yielding leptoceratopsid remains (slightly modified from fig. 1 in Lindgren and Siverson 2002).

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Figure TEXT-FIG. 3..   Leptoceratopsidae gen. et sp. indet.; selected isolated teeth from the lowermost Campanian Gonioteuthis granulataquadrata Zone (RM PZ R1833–1834) and the informal, uppermost Lower Campanian Belemnellocamax mammillatus zone of the Kristianstad Basin; × 3. A–B, RM PZ R1827, a right maxillary tooth in distal and mesial views (stereo pairs). C–E, RM PZ R1828, a right maxillary tooth in C, distal, D, occlusal and E, mesial views. F–I, RM PZ R1833, a right maxillary tooth in F, mesial, G, labial, H, lingual and I, distal views. J–K, RM PZ R1830, a left dentary tooth in mesial and distal views (stereo pairs). L–N, RM PZ R1834, a right dentary tooth in L, distal, M, lingual and N, mesial views.

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Figure TEXT-FIG. 2..   Line drawings of A, RM PZ R1833 and B, RM PZ R1830 in distal and mesial views, respectively, showing the dental terminology used in the text.

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A discontinuity surface with burrows marks the Lower/Upper Campanian boundary at this locality. A lag deposit of reworked Belemnellocamax mammillatus (Nilsson, 1826) is present at the base of the Upper Campanian (Lindgren and Siverson 2002). Primitive Belemnellocamax balsvikensis (Brotzen, 1960) (an index taxon for the lowermost Upper Campanian in southern Sweden; Christensen 1975) appear a few centimetres higher up.

Ullstorp 1 sensuErlström and Gabrielson (1986) (Text-fig. 1). Map sheet: Kristianstad 3D SO (topographical map sheet, 1 : 50,000); coordinates UTM VC 165 849 (56°03′46·2″N, 13°57′21·0″E). Christensen (1975) described the belemnite fauna from the calcarenites in the Ullstorp quarries and assigned all exposed strata (including those in Ullstorp 3, which is referred to as Ullstorp 1 in Erlström and Gabrielson 1986) to the informal Belemnellocamax mammillatus zone of the uppermost Lower Campanian based on the occurrence of Belemnellocamax mammillatus, Belemnitella mucronata (von Schlottheim, 1813) and Gonioteuthis quadrata scaniensisChristensen, 1975.

Continued operation in one of the quarries in the 1980s exposed a 10-m-thick sequence of terrigenous clastics underlying uppermost Lower Campanian biocalcarenites. This prompted Erlström and Gabrielson (1986) to investigate the quarry and to publish a description of the sediment and its fauna. They referred the entire terrigenous clastic sequence to the latest early Campanian, based on the discovery of Belemnellocamax mammillatus in the basal part of the section.

Extensive collecting in two of the conglomerates (beds C and E of Erlström and Gabrielson 1986, fig. 3) in the predominantly siliciclastic sequence over the last two decades has produced a large number of vertebrate remains (including two leptoceratopsid teeth) in addition to numerous belemnites (Siverson 1993b). We have not found any of the belemnite species characteristic of the informal Belemnellocamax mammillatus zone and can therefore conclude that Erlström and Gabrielson's (1986) age assignment for the terrigenous clastic section at Ullstorp 1 is incorrect. Their ‘Belemnellocamax mammillatus’ specimen from the basal part of the section (J. Gabrielson, pers. comm. 2004) is almost certainly a Belemnellocamax grossouvrei (Janet, 1891), a species relatively common in both conglomerate C (juvenile to subadult specimens) and E (mostly adult specimens). This species along with Actinocamax verusMiller, 1823 and Gonioteuthis granulataquadrata (Stolley, 1897) in conglomerate C places the bed in the lowermost Lower Campanian Gonioteuthis granulataquadrata Zone.

Institutional abbreviations.  AMNH, American Museum of Natural History, Division of Paleontology, New York, USA; CMN, Canadian Museum of Nature, Ottawa, Canada; RM PZ, Swedish Museum of Natural History, Department of Palaeozoology, Stockholm, Sweden; TMP, Royal Tyrrell Museum of Palaeontology, Drumheller, Canada.

Tooth terminology.  In accordance with the recommendations of Smith and Dodson (2003), the following descriptive terms are used herein: mesial (anterior), distal (posterior), lingual (towards the tongue), labial (towards the lip), apical (towards the tip of the cusp) and basal (towards the root).


Remarks.  Traditionally (Brown 1914; Sternberg 1951; Dodson and Currie 1990) all small-bodied, basal neoceratopsians were lumped together in the Protoceratopsidae. However, recent cladistic analyses have unanimously shown this family to be paraphyletic, and thus of somewhat limited taxonomic value (Sereno 1986, 2000; Makovicky 2001; Xu et al. 2002; You and Dodson 2003). There are currently a small number of loosely defined subgroups of basal neoceratopsians supported by one or more synapomorphies (Sereno 2000).

The dental apparatus of most neoceratopsians comprises maxillary and mandibular teeth, while the rostral end of the skull terminates in a beak-like snout. Premaxillary teeth are present in a few primitive forms, such as Archaeoceratops and Liaoceratops (Xu et al. 2002; You and Dodson 2003), but were apparently lost during the evolution of the higher Neoceratopsia (Sereno 1986). Isolated teeth of leptoceratopsids are readily distinguished from those of other ceratopsids on the basis of the following character state combination (Ostrom 1978; Baszio 1997; Sereno 2000): (1) root large but not bifurcated and broader transversally than it is mesiodistally long; (2) newly erupted teeth have ovate crowns in labial and lingual views and are broader labiolingually than mesiodistally; (3) crowns of functional teeth unilaterally enamelled (but see also discussion below); (4) each tooth has a strong lingual (dentary) or labial (maxilla) primary ridge that merges with a marked basal crown cingulum; (5) secondary ridges prominent and either subparallel or obliquely inclined to asymmetrically placed primary ridge; and (6) occlusal wear surfaces on dentary teeth horizontally ledged.

LEPTOCERATOPSIDAE gen. et sp. indet. Text-figures 2–4


Figure TEXT-FIG. 4..   Leptoceratopsidae gen. et sp. indet., caudal vertebral centra from the informal, uppermost Lower Campanian Belemnellocamax mammillatus zone of the Kristianstad Basin; × 1·1. A–C, RM PZ R1831, in A, ventral (stereo pair), B, lateral and C, rostral views; note the asymmetrical, possibly pathological morphology of the rostral surface. D–E, RM PZ R1832 in ventral and lateral views (stereo pairs).

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Material.  Åsen, three maxillary teeth (RM PZ R1827–1829), one dentary tooth (RM PZ R1830) and two caudal vertebrae (RM PZ R1831–1832). Ullstorp, one maxillary tooth (RM PZ R1833) and one dentary tooth (RM PZ R1834).


Maxillary teeth.  The maxillary teeth from the Kristianstad Basin leptoceratopsid are moderately (Text-fig. 3F–I) to strongly (Text-fig. 3A–B) mesiodistally flattened with an elliptical (Text-fig. 3D) to almost rhomboidal occlusal outline. A thick layer of enamel is present on the occlusal and labial sides of the crown, whereas the lingual surface comprises two inclined wear facets. A conspicuous, lingually worn cusp occupies the medial half of the occlusal surface, whereas the labial half of the crown is developed into a roughly horseshoe-shaped horizontal shelf. In mesial and distal views, the two surfaces are set nearly perpendicular to each other (Text-fig. 3A–B). The labial surface of the cusp is asymmetrically divided into a wide, mesiolaterally sloping mesial indentation and a narrow distal plane by a primary ridge. This ridge and a small number of secondary ridges are ornamented with numerous small and densely packed bumps. In addition to these bumps, a few faint lines are present on the mesial face of the anteriormost secondary ridge (Text-fig. 3B). The enamel cover is intact at the centre of the labial shelf, whereas it is damaged along the edges of this surface, forming a distinct V-shaped occlusal wear facet (Text-fig. 3D). Judging from the partially preserved bony base in RM PZ R1833 (Text-fig. 3F–I), the single root of each tooth was originally long and transversely broad. Moreover, the root of RM PZ R1833 is strongly mesiodistally compressed, and shallow, longitudinal grooves on either side contribute to its somewhat hour-glass-shaped horizontal outline. As preserved, the largest maxillary tooth (RM PZ R1827; Text-fig. 3A–B) measures 10·2 mm in maximum width and 5·4 mm in total length (at the base of the enamel).

Dentary teeth.  With a more or less convex lingual side and a concave labial face, each of the leptoceratopsid dentary teeth from the Kristianstad Basin has an almost ‘claw-like’ mesial profile (Text-fig. 3J, N). The apex of the ‘claw’ is missing, however, and instead there are two inclined occlusal wear facets. In RM PZ R1830 (the best preserved of the two dentary teeth), the inner facet slopes at c. 40 degrees to the vertical axis of the tooth (measured perpendicular to the plane formed by the basal crown cingulum), whereas the outer one forms a well-defined ledge (Text-fig. 3J–K). The angle between the two facets is close to 120 degrees. A longitudinal, somewhat labially curved groove reaches almost to the base of the crown on the mesial side (Text-fig. 3J). It is bordered lingually by a thickly enamelled secondary ridge, whereas the floor of the depression itself is seemingly devoid of enamel. A prominent primary ridge divides the lingual surface into two enamel-covered indentations (the distal indentation has approximately three times the area of the mesial one). Two secondary ridges (one in front and one behind the primary ridge) are subparallel to the primary ridge and constitute the most distal and mesial parts of the crown, respectively. The base of the lingual surface of the crown has a narrow cingulum that partly embraces small pockets on either side of the primary ridge (Text-fig. 3J). As preserved, the enamelled labial face is markedly lower than the lingual face and rather narrow mesiodistally. A minutely ribbed and almost vertical protuberance is posteriorly directed on the distal face of the crown (Text-fig. 3K). The outgrowth is enamelled at the base, but lacks enamel at its apex. Only a vestige of the root is preserved in RM PZ R1830. It can, nonetheless, be concluded that the transition from the crown to the root is gradual: there is no great size discrepancy in any dimension between the base of the crown and the top of the root. The largest tooth at hand, RM PZ R1834 (Text-fig. 3L–N), is 8·6 mm wide (measured at the base of the enamel), 5·7 mm long (mesiodistally) and 8·9 mm high (the vertical distance from the base to the tip of the crown disregarding its curvature).

Vertebrae.  Two caudal vertebrae, RM PZ R1831 (Text-fig. 4A–C) and R1832 (Text-fig. 4D–E), from the local, uppermost Lower Campanian Belemnellocamax mammillatus zone are here tentatively referred to the Kristianstad Basin leptoceratopsid. The morphology is consistent with the caudal anatomy of basal neoceratopsians, and the tentative identification is parsimonious considering the presence of leptoceratopsid teeth in the same deposits. The interarticular surfaces of at least RM PZ R1831 appear to have been rounded in life (Text-fig. 4C), and both centra display caudally inclined ventral facets for articulation with chevrons of the haemal arch. As preserved, RM PZ R1831 is 25·5 mm long and 34·6 mm wide (original width is estimated at c. 37 mm), whereas RM PZ R1832 measures 25·8 mm in total length and 27·2 mm in maximum width.

Comparisons.  Teeth of neoceratopsians are unilaterally enamelled, with the distribution of the enamel on opposite sides of upper and lower crowns (Galton 1973, fig. 5C–D; Baszio 1997; Sereno 2000). By contrast, dentary teeth of the Kristianstad Basin leptoceratopsid seem to have enamel on both sides of the crown, although the enamel cover is thicker on the lingual face than on the labial side. Ledged occlusal wear facets were considered a diagnostic feature of Leptoceratops gracilisBrown, 1914 by Ostrom (1978), although a similar pattern of wear has subsequently been found on teeth of MontanoceratopsSternberg, 1951,UdanoceratopsKurzanov, 1992 (see Chinnery and Weishampel 1998) and now the Swedish neoceratopsian.

Although the neoceratopsian maxillary and dentary teeth from the Kristianstad Basin could theoretically originate from different species, their matching size, overlapping gross crown morphology, and the fact that the wear facets on the maxillary teeth would seem to fit into the occlusal notch on the dentary teeth strongly suggest that all specimens represent the same taxon. This neoceratopsian differs from all other described basal neoceratopsians, including LeptoceratopsBrown, 1914 and Udanoceratops (Brown 1914, fig. 6; Kurzanov 1992, fig. 4C–D; pers. obs. of AMNH 5205, CMN 8889), by its markedly mesiodistally flattened maxillary crowns and by the wide labial shelves of the maxillary teeth. The overall morphology and occlusal wear configuration of the Swedish dentary teeth correspond roughly to those of Leptoceratops (Ostrom 1978, pl. 2, figs 2–3), although the crowns from the Kristianstad Basin lack the numerous, densely spaced secondary ridges that are seemingly typical of the North American genus (Ryan and Currie 1998, figs 3C, 4). Even though the neoceratopsian from the Kristianstad Basin probably represents a new taxon (considering that the Fennoscandian Shield was an isolated land mass throughout the Late Cretaceous), the fragmentary nature of the known specimens does not allow a meaningful diagnosis. There are, for instance, no complete and unworn teeth in the leptoceratopsid collection from the basin. Therefore, a genus and species has not been assigned to the Swedish leptoceratopsid, pending the discovery of more complete material from the Upper Cretaceous of the basin.

Remarks.  Whereas the occlusion in all basal neoceratopsians and higher ceratopsians is exclusively one of shear (Brown and Schlaikjer 1942, fig. 21B; Ostrom 1964, 1966; Galton 1973), the dentition of Leptoceratops, Montanoceratops and Udanoceratops combines the functions of cutting and crushing (Sternberg 1951; Ostrom 1966, fig. 4; Galton 1973, fig. 5E; Chinnery and Weishampel 1998). This specialization probably enabled these animals to chop their food into small pieces using their lingual shearing surfaces and then to crush it in the labial notch of the dentary teeth. The enlarged labial shelf of the dentary teeth (Text-fig. 3J) in combination with the wide crushing surface on the maxillary teeth (Text-fig. 3A) would have provided the Kristianstad Basin leptoceratopsid with a masticatory apparatus similar to those of Leptoceratops, Montanoceratops and Udanoceratops.

It has been suggested (Galton 1973) that the broad and shelf-like mandibles in Leptoceratops were used to collect chewed plant litter. If correctly interpreted, it is reasonable to assume that the Kristianstad Basin leptoceratopsid was also equipped with large lateral dentary shelves to catch broken-down plant debris. Furthermore, the distinctly mesiodistally flattened and laterally grooved roots suggest that the teeth of the Swedish neoceratopsian were packed in some form of magazine (tooth batteries). However, contrary to the complex tooth batteries of the more advanced ceratopsians, in which the transversely bifurcated roots straddle a number of succeeding replacement teeth at any given time (Ostrom 1964), the single root in the Kristianstad Basin leptoceratopsid must have limited the number of replacement teeth to one or two. Presumably, marginal portions of the crowns of adjacent germ teeth could fit into the shallow depressions on the mesial and distal surfaces of the root in a fashion comparable with that of protoceratopsids (Brown and Schlaikjer 1942).

Material and methods

  1. Top of page
  2. Abstract
  3. Geological setting and localities
  4. Material and methods
  5. Systematic palaeontology
  6. References

The material was collected by sifting the poorly consolidated sands at Åsen and the more indurated conglomerate at Ullstorp through sieves with a mesh width of about 2·5 mm. Adhering carbonates were removed by treatment with buffered acetic acid (Jeppsson et al. 1999), and the fossils were hardened by application of polyvinylbutyral in acetone. Illustrated specimens were painted with graphite and coated with ammonium chloride prior to being photographed. For descriptive terms used in the text, see Text-figure 2.

Systematic palaeontology

  1. Top of page
  2. Abstract
  3. Geological setting and localities
  4. Material and methods
  5. Systematic palaeontology
  6. References


On a broad biogeographical level, the Ceratopsia has hitherto had a bimodal distribution pattern in the Northern Hemisphere with occurrences restricted to central Asia and western North America (Sereno 2000). A central Asian origin for the group has been generally accepted, and, until recently, the predominant view was that ceratopsians arrived in North America following one or more unidirectional dispersal events from Asia across the periodically open Bering land bridge (Maryańska and Osmólska 1975; Chinnery and Weishampel 1998; but see also Chinnery et al. 1998). However, Sereno (1997, 1999, 2000) demonstrated that the biogeographical history of the Ceratopsia is much more complicated than previously thought, and that the phylogeny of the clade requires multiple, bidirectional dispersal events between Asia and North America. Most dispersal scenarios favour a polar sweepstakes dispersal route across Beringia (Sereno 2000), although Chinnery et al. (1998) suggested an alternative pathway across ancient Europe. This latter option has been largely rejected, however, owing to the lack of ceratopsian remains from Europe. The occurrence of leptoceratopsid teeth and bones in the Kristianstad Basin changes the prerequisites for a European dispersal route and may necessitate a new interpretation of neoceratopsian biogeography.

As defined by Makovicky (2001), the Leptoceratopsidae (sensu stricto) comprises Leptoceratops and Montanoceratops, and is, hence, known solely from the Campanian and Maastrichtian of the USA and Canada (Ryan and Currie 1998, table 1; You and Dodson 2003). Recently, Chinnery (2004) described Prenoceratops pieganensis from Montana. However, a fourth possible leptoceratopsid (or a very closely related genus), Udanoceratops, was described by Kurzanov (1992) from Santonian–Middle Campanian deposits of Udan-Sayr in southern Mongolia, thus widening the biogeographical range of leptoceratopsid-like neoceratopsians beyond North America. The new leptoceratopsid record from the Lower Campanian of the Kristianstad Basin may, on the one hand, support a European dispersal route for the Leptoceratopsidae, as it connects the Asian and North American leptoceratopsid and leptoceratopsid-like faunas. In a European (Scandinavian) dispersal scenario, the group would have arrived in eastern North America first, after dispersing across Scandinavia from Asia, hence supporting the hypothesis of Chinnery et al. (1998) of a mid-Cretaceous faunal migration to North America from Eurasia.

On the other hand, the Kristianstad Basin population of leptoceratopsids may represent an entirely endemic assemblage of horned dinosaurs. Considering that the Fennoscandian Shield was an isolated land mass throughout the Late Cretaceous (Smith et al. 1994), this latter possibility seems to be the most likely one. A land bridge connecting west-central Asia and the Fennoscandian Shield was probably present during late Early Cretaceous times (Smith et al. 1994, map 16), and the presence of basal ceratopsians in central Asia in strata that pre-date the earliest Early Campanian Gonioteuthis granulataquadrata Zone of the Kristianstad Basin (the Ullstorp locality) indicates that (in the absence of other known occurrences of ceratopsians outside of Asia, North America and Sweden) the Swedish population of leptoceratopsids descended from individuals that migrated westwards from west-central Asia rather than vice versa. Isolation commenced during the early Late Cretaceous when the Turgai Strait reached the northern part of the Tethyan Ocean, thus preventing passage between Asia and Scandinavia (Smith et al. 1994, map. 15).

It has long been assumed that basal neoceratopsians, including leptoceratopsids, preferred arid and/or semi-arid habitats, and only occasionally entered coastal-plain environments (Sternberg 1951; Ryan and Currie 1998; but see also Ott and Buckley 2001). This supposition is based largely on the frequent occurrence of articulated protoceratopsian skeletons in the aeolian deposits of central Asia as opposed to the relative scarcity of leptoceratopsids in the predominantly nearshore Cretaceous strata of western North America. Nonetheless, the leptoceratopsid remains from the Kristianstad Basin were collected from shallow, nearshore marine strata deposited in an archipelago setting with rocky shorelines and low islands (Jensen 2002). Although the teeth and vertebrae may originate from drifting and bloated carcasses that were swept out to sea from inland areas by rivers during flood events, it is likely that the animals to which the fossils originally belonged preferred coastal habitats and may even have inhabited larger islands in the archipelago. For instance, the neoceratopsian teeth from the Ullstorp site were found in a calcareous conglomerate (conglomerate C sensuErlström and Gabrielson 1986) that accumulated in a nearshore environment within the storm wave base to breaker zone interval. At the time of deposition, this locality was situated a few hundred metres to the north of the Nävlingeåsen ridge (presumably an island during the Early Campanian), whereas the coherent Fennoscandian land mass was located several tens of kilometres further to the north (see the palaeogeographical reconstruction in Jensen 2002, p. 24).

Acknowledgements.  We gratefully acknowledge logistical support for the project provided by the Department of Geology, GeoBiosphere Science Centre, Lund University. Three successful field seasons at Åsen and Ullstorp were made possible through financial support from Crafoordska stiftelsen, Längmanska kulturfonden and Kungliga Fysiografiska Sällskapet i Lund. Margaret Feuerstack (Canadian Museum of Nature) and Carl Mehling (American Museum of Natural History) provided photographs of CMN 8889 and AMNH 5205 (Leptoceratops gracilis), respectively. Reviews by Michael Ryan and an anonymous referee greatly improved the final version. To all our sincere thanks.


  1. Top of page
  2. Abstract
  3. Geological setting and localities
  4. Material and methods
  5. Systematic palaeontology
  6. References
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