Late Cretaceous (Campanian) leaf and palynoflora from southern Skåne, Sweden

Abstract A Late Cretaceous (Campanian) leaf megaflora from the Vomb Trough in southern Skåne, Sweden, has been investigated on the basis of collections held at the Swedish Museum of Natural History. The main plant-bearing locality is Köpinge, but single specimens originate from Högestad, Ingelstorp, Rödmölla, Svenstorps mölla and Tosterup. The fossil flora is dominated by the angiosperm (eudicot) Debeya (Dewalquea) haldemiana (Debey ex de Saporta & Marion) Halamski. Other dicots are cf. Dryophyllum sp., Ettingshausenia sp., Rarytkinia? sp., Dicotylophyllum friesii (Nilsson) comb. nov. and Salicites wahlbergii (Nilsson) Hisinger. Conifers are represented by cf. Aachenia sp. (cone scales), Pagiophyllum sp. and Cyparissidium sp. (leaves). Single poorly preserved specimens of ferns and monocots have also been identified. The terrestrial palynomorphs (the focus herein) clearly link to the megaflora, although with different relative abundances. The fern spore Cyathidites dominates along with the conifer pollen Perinopollenites elatoides and Classopollis. Angiosperm pollen comprise up to 15% of the assemblage, represented by monocolpate, tricolpate and periporate pollen and the extinct Normapolles group. The spores in the kerogen residue show a thermal alteration index (TAI) of 2+. The flora probably represents mainly a coastal lowland Debeya/conifer forest, and is similar to approximately coeval assemblages from analogous palaeo-communities described from eastern Poland, western Ukraine and Westphalia.

Late Cretaceous terrestrial to shallow-marine rocks of Skåne, southern Sweden yield plant micro-, meso-and megafossils. The micro-and mesoflora have been the subject of several studies over the past three decades (mostly from northern Skåne: see the review by Friis et al. (2011, pp. 70-72)). The wood remains and fossil cones were described by Conwentz (1892), Nykvist (1957), Srinivasan & Friis (1989) and Herendeen (1991). However, the leaf megaflora has only been reported and briefly described in a few publications from the first half of the nineteenth century (Nilsson 1824(Nilsson , 1832Hisinger 1837), and no palynological analyses have been performed on the Campanian successions from the Vomb Trough. This paper aims to revise the Campanian leaf flora from the Vomb Trough in southern Skåne (Fig. 1) and to compare the megaflora to the palynological assemblage recovered from the same strata. Santonian leaves and cones from the Kristianstad Basin in northern Skåne will be dealt with in a separate study.

History of research
Fossil plants from the Late Cretaceous deposits of Skåne were first mentioned and illustrated by Nilsson (1824). Sven Nilsson (1787Nilsson ( -1883 began collecting objects of natural history in his youth, becoming Associate Professor in Natural History at Lund University by 1812. He took up the directorship of the Swedish Museum of Natural History in Stockholm (NRM) in 1828, but in 1832 he returned to Lund taking up a professorship in natural history until 1856. During his time at NRM, he assembled a collection that served as the basis for the first detailed study of Swedish Cretaceous plants, with the description of five new species from Köpinge (Nilsson 1832). Nilsson's palaeozoological collections were donated to the museum in Lund and have been used by subsequent researchers (e.g. Lundgren 1865Lundgren , p. 1, 1874Lundgren , p. 70, 1880). Among the palaeobotanical collections available to the present authors, only two specimens (S084165 and Lund s.n. (s.n., sine numero)), however, can be attributed unambiguously to the Nilsson collection. Some of his plant fossils are, perhaps, present amongst a collection of material originating from Lund but now (2015) registered at the NRM, but the documents detailing the collector(s) and transfer are lacking. This concerns, in particular, type and figured specimens that could not be traced for the purpose of the  Chatziemmanouil (1982), Surlyk in Voigt et al. (2008) and Halamski (2013)), and the presumed location of the studied flora and of three other coeval palaeofloras (circle, Poland-Ukraine; square, Haldem in Westphalia; triangle, Grünbach in Austria). (b) Bedrock geological map of Skåne (Scania) and adjacent regions after Koistinen et al. (2001), Vajda & Gravesen (2008) and references therein. (c) Solid geological map of the Vomb Trough in southern Skåne showing the localities that have yielded the fossil flora studied, after Christensen (1986). present study. Nilsson's specific descriptions were repeated by Hisinger (1837), who moreover introduced four new genera based on these species. He later relisted the four dicotyledonous species from Köpinge in a synoptic table of fossils from Skåne (Hisinger 1840, p. 58).
Another major collection from the studied area (available to the present authors) was assembled by a collector named Gustaf von Schmalensée (1842-1919, and was deposited in the NRM during the years 1879, 1880 and 1903. It partly derives from the Swedish Geological Survey (Sveriges Geologiska Undersökning) collections, as testified by a letter of Edvard Erdmann to Alfred Gabriel Nathorst, a professor at the NRM, dated 5 November 1902. Nathorst (1881) had already published a short nomenclatural remark on the most abundant species at Köpinge and apparently planned to undertake a more extensive study of this flora, based on the names attributed to him occurring on collection labels. However, these names have never been published.
'Dewalquea nilssonii' was listed among fossils from Köpinge by Moberg (1884). The flora of 'Köpinge Sandstone' was then briefly mentioned by Hartz (1909, p. 2), who described it as consisting of: 'fragmentary rests of ferns, conifers, and angiosperm trees; the most common leaf being that of Dewalquea Nilssonii (Helleboreae?)'. A similarly laconic remark was made by Ross (1949, p. 38), who stated: 'a few dubious dicotyledonous fossils [Acerites, Alnites, Cycadites (Dewalquea), Salicites] from the Upper Cretaceous of Scania'. Unlike the floras from the Kristianstad Basin, Cretaceous plants from southern Skåne have not been studied for more than a century.

Geological setting
Skåne (also known as Scania) is a province of southern Sweden that incorporates the geological transition (Sorgenfrei-Tornquist Zone) between the Baltic (Fennoscandian) Shield and the Danish Basin within the Northwest European Craton. Mesozoic strata are known in Skåne from both outcrops and borings. Cretaceous deposits are known from several areas (Fig. 1b), the most important from a palaeobotanical perspective being the Kristianstad Basin, situated on the southernmost part of the Fennoscandian Shield, and the Vomb Trough, situated in the Fennoscandian Border Zone (Christensen 1984;Larsson et al. 2000). The Late Cretaceous landscape of Skåne consisted of several elongate islands and peninsulas (Fig. 1a) formed of basement (Precambrian-Palaeozoic) rocks surrounded by shallow-marine areas (Surlyk in Voigt et al. 2008, p. 940). Å sen, a well-known fossil site within the Kristianstad Basin, hosts charcoalified flowers within Santonian-lower Campanian fluvial deposits (Skarby 1968;Friis 1984;Friis et al. 2011). Within the Kristianstad Basin, the fluvial successions are overlain by marine glauconitic calcareous sandstones hosting a rich fossil biota, including a range of invertebrates (Einarsson et al. 2016), but also vertebrates including sharks and fish Siversson et al. 2015), mosasaurs, plesiosaurs (Einarsson et al. 2010) and even non-avian dinosaurians (Poropat et al. 2015). Oysters were described from Å sen showing heteromorphic ornamentation interpreted as attachment in vivo to arborescent plants (presumably mangroves: Friis & Skarby (1981)).
The Vomb Trough (Fig. 1c) is a graben extending for about 55 km northwestwards and 25 km southeastwards from Ystad (Norling 1982). It is bordered by the Precambrian Romeleåsen Horst to the SW and by a lower Palaeozoic plateau to the NE: both were also areas of elevation in the Cretaceous (Chatziemmanouil 1982) and, thus, may be assumed to be the source of the flora described from the localities situated in that region. The Upper Cretaceous stratigraphy of the Vomb Trough was elaborated by Christensen (1986) on the basis of belemnites.
The richest Late Cretaceous megafloral assemblage in Skåne derives from Köpinge. The name of this locality (also Köpinge mölla) situated approximately 10 km NE of Ystad (not to be confused with Köpinge 10 km SE from Kristianstad) denotes a large number of small outcrops (see detailed description by Hägg (1954, pp. 11 -13): most of them described as already inaccessible by Christensen (1986)) of a yellow calcareous sandstone rich in glauconite, the so-called 'Köpinge sandstone'. Besides plants, the 'Köpinge sandstone' has yielded foraminifera (Brotzen 1938), molluscs (Schlüter 1870), brachiopods (Dalman 1828;Hägg 1954) and vertebrates (Nilsson (1835): see the list published by Moberg (1884, p. 35) and the bibliography in Christensen (1986, p. 9)). The rock is sufficiently characteristic to also be recognized among erratic boulders (Foth et al. 2011). The age of the 'Köpinge sandstone' (Christensen 1986) is latest early Campanian and middle late Campanian (strata of earliest late Campanian age are apparently absent). The locality Svenstorp (or Svenstorps mölla) is situated in the greater Köpinge area (Hägg 1954, pp. 11 -12). Ingelstorp is situated immediately SE of Köpinge and the sandstone exposed in this parish is about the same age as that from the latter locality (Hägg 1954, p. 15). Högestad is situated about 10 km NW of Köpinge and strata exposed there belong to the middle part of the Campanian (Mucronatenkreide sensu Hägg (1954, pp. 11, 15); see also Christensen (1976, pp. 16-17)).
The localities Rödmölla and Tosterup, situated approximately 2 km east of Köpinge, require a slightly more detailed comment. A series of outcrops formerly existing in that area exposed several Upper Cretaceous beds (Christensen 1976, pp. 11-12; see also Moberg 1884, pp. 31-34). Some of them belong to strata older than the succession at Köpinge (Mamillatenkreide sensu Hägg 1954, p. 4); however, it may be assumed that the plant material comes from the Tosterup conglomerate dated to the latest early Campanian on the basis of belemnites (Christensen 1976, p. 13).

Megaflora
The studied material consists of 160 specimens, 147 of them (92%) derived from Köpinge. The absolute and relative abundances of the various taxa identified in this collection are summarized in Table 1. Plant fossils were studied entirely from historical collections, as the outcrops are long since inaccessible. Most of the name-bearing specimens (types) of Nilsson (1832) could not be found and are probably lost. This is particularly unfortunate because taxonomic interpretations of the relevant taxa will necessarily be hypothetical. Taxa that could not be identified due to a lack of specimens are Acerites cretaceus (Nilsson, 1832) Hisinger, 1837, Comptonites antiquus (Nilsson, 1832) Hisinger, 1837 and Cannophyllites septentrionalis Nilsson, 1832. The studied collection of 159 specimens is housed in the NRM in Stockholm. A single specimen is housed in the type collection at the Department of Geology, Lund University, Sweden.
The plant fossils are fragmentary or incomplete leaves preserved as imprints in calcareous sandstone (Figs 2-7) and many are curved (i.e. not preserved along a single plane): this suggests high-energy sedimentation or bioturbation. The leaves are distinguished from the surrounding rock owing to their brownish coloration (Fig. 2a, c -f ), likely to be a result of bacterially mediated ferric-oxide   (e) cf. Aachenia sp., a conifer cone scale, specimen S084113, Köpinge; (f ) subcomplete compound leaf S166107 of Debeya (Dewalquea) haldemiana; and (g) fragmentary leaflets of Debeya (Dewalquea) haldemiana, the arrangement of which suggests that they come from a single compound leaf, specimen S084187, Köpinge. (c) twig S084212, Rödmölla; (d) twig S166168, Köpinge; (e) twig S166108, Köpinge; (f ) twig S083929, Köpinge; (g) twig S166150 cut transversely, Köpinge. precipitation. Such preservation is rather similar to that of the coeval flora from eastern Poland and western Ukraine (Halamski 2013). Rare specimens are blackish (Figs 2b & 5c), suggesting a different type of preservation, but this has not been investigated in detail. Concerning the venation, in all leaves belonging to Ettingshausenia sp. tertiaries and quaternaries are preserved, and in Debeya haldemiana, two specimens bear faint traces of tertiaries (Fig. 5h), whereas in all other leaves only the midvein and the secondaries can be observed. Overall, the preservational state of the megaflora may be described as rather poor. Uncoated specimens were photographed under oblique and most often polarized light, using a Canon 350D digital camera either with a Canon EFS 17 -85 mm lens or, if required, with a Canon EF 100 mm macro lens.

Palynology
Two rock samples from the Köpinge locality (NRM-S089754 and NRM-S089755), each yielding leaf macrofossils, were selected for palynological analysis. Around 10 g of beige, fine-grained sandstone were processed according to standard palynological procedures at Global Geolab Ltd, Canada (e.g. see Vajda et al. 2013). The organic residue was sieved using a 5 mm mesh and mounted in epoxy resin on two microscopic slides. Three hundred pollen and spores were identified per sample and the percentage of each palynomorph taxon was calculated (Table 2) In addition, one kerogen slide was prepared from each sample (i.e. where the residue did not go through the oxidation or sieving processes) in order to estimate the thermal alteration index (TAI) of the palynomorphs.
Palynofacies analysis involved counting the relative abundance of organic particles based on 500 counts per slide. Classification of the palynological matter was based on the scheme developed by Batten (1996), and the groups identified were: (I) palynomorphs (pollen, spores, dinoflagellates and algae); (II) wood; (III) charcoal; (IV) plant cuticle; and (V) amorphous organic matter (AOM). The slides and residues are deposited in the palaeobiology collections of the NRM, Stockholm, and light micrographs of selected pollen and spores from the two studied samples are illustrated in Figure 8 and are identified by NRM numbers and Englandfinder coordinates.

Systematic palaeontology (megaflora)
Infrakingdom Cormophyta (Endlicher, 1836) Cavalier   identical) to those reported as Microphyllopteris by Halamski (2013 Description. Ovuliferous cone scale 14 mm long and 12 mm wide, triangular in outline, with an attenuate apex and an inconspicuous keel. The basal region of the scale bears three impressed depressions  probably representing seeds attached at the base of the scale. The seeds are ovoid, 4-5 mm long and 2-3 mm wide. It is unclear whether the lack of a basal stalk characteristic of Aachenia is a preservation artefact or if the stalk was originally missing. Remarks. This ovuliferous scale is broadly similar to scales described as Aachenia debeyi from the Late Cretaceous of Aachen (Knobloch 1972) and A. knoblochii Wagner from the Maastrichtian of Mexico (Weber 1985 fig. 7). Rödmölla: S084212.
Remarks. Leaves are longer than wide, a characteristic feature of Pagiophyllum as reintroduced by Harris (1969 Remarks. We employ this genus in the sense of its emendation by Harris (1969, pp. 248-249; see also Harris 1979, p. 77).
Description. The single available specimen represents a fragment of a leafy twig 23 mm long with numerous, dense, helically arranged, needle-like leaves. The leaves are 0.7 × 3 mm and appressed to the axis, forming a branch 3 mm in diameter.
Remarks. Debeya haldemiana is by far the most common plant species in the material studied. The specimens are usually leaflets showing no organic connection, only rarely do they represent incomplete leaves (apical parts always lacking). In comparison to other localities from which this species was reported (Haldem, Potylicz), the striking feature of the Köpinge material is the common occurrence of small leaves (up to 4 mm in width compared with presumably adult leaves 10 mm or more wide).
Remarks. This specimen is compared to Dryophyllum (as emended by  on account of its elliptical shape, pinnate craspedodromous venation and asymmetry of the secondaries, suggesting that it is, in fact, a leaflet of a pinnately compound leaf. However, interpretation of the single poorly preserved specimen is equivocal at best. Ettingshausenia sp. (Fig. 6b-e) Material. Köpinge: S084034.
Description. Leaves entire-margined or with few teeth, trilobate with lobes poorly developed. The fragments vary in size, the smallest being 40 × 20 mm and the largest over 50 mm wide, suggesting that complete leaves were relatively small (c. 7 cm in width and length). Leaf lamina obovate. Apex acute (Fig. 6e); base probably cuneate as fragmentarily preserved (Fig. 6c). Lobes very small (Fig. 6e) or nearly absent (specimen S084207). Petiole preserved in a single specimen (Fig. 6d). Venation palmately pinnate semibrochidodromous. Midvein stout, terminating in the leaf apex. Two pairs of suprabasal secondary veins emerging obliquely from the midvein. No infrabasal secondary vein. Secondaries semibrochidodromous (Fig.  6e). Tertiary veins prominent, straight or slightly curved, percurrent.
Remarks. The studied material is very similar in general venation style and size to Ettingshausenia gruenbachiana Herman & Kvaček, 2010 from the lower Campanian of Grünbach. It differs in its more angular leaf form and less pronounced lobes. The studied material differs from Ettingshausenia? lublinensis (Karczmarz & Popiel, 1971 Remarks. This leaf is tentatively ascribed to Rarytkinia on the basis of its large size and the departure angle of the secondary veins being significantly high in the basal region, but low and constant in the rest of the leaf. A coeval representative of the genus is the insufficiently known Rarytkinia polonica (Karczmarz & Popiel) Halamski from eastern Poland (Halamski 2013), which differs from the form described here in its wider shape and denser secondaries. Some leaves in the collection are labelled 'Protophyllum suecicum Nath. mscr.', but this name has never been published and, thus, has no nomenclatural status; the available material is insufficient for a formal description.

Form group unknown
Genus Dicotylophyllum auct.

Remarks. Alnites Hisinger is a synonym of Alnites
Dicotylophyllum friesii (Nilsson, 1832) comb. nov. (Fig. 7a- Material. Köpinge: S083980. Description. The available material consists of leaf fragments, one representing the basal and median regions of the lamina, and the other the apical region: the base and apex themselves, however, were not preserved. Leaf elliptical, estimated length of approximately 8-10 cm, estimated width approximately 2 cm. Midvein straight. Secondary veins partly decurrent and partly excurrent (Fig.  7e), alternate, angle decreasing from c. 608 in the basal region through to c. 408 in the middle of the lamina to c. 308 in the apical region; spacing irregular. Margin serrate with one order of teeth and three teeth per cm; teeth short, proximal side convex, distal side weakly concave.
Remarks. In the absence of the holotype, the interpretation of the simple description and somewhat schematic figure given by Nilsson (1832) is not easy. However, the finely serrate ('subcrenatus') margin and the elliptical shape make the conspecificity of the material described here with Nilsson's species probable. A neotype is selected to fix the usage of the species name.
Remarks. Again, the interpretation of Nilsson's species in the absence of types is not easy. However, the absence of teeth and the irregular spacing of secondary veins is consistent with the figure in the protologue.
Description. The single available specimen consists of two leaf fragments on a single rock sample, one fragment is approximately 17 mm long and 5.5 mm wide, and the other is approximately 15 mm long and 8 mm wide. The margins of the smaller fragment run parallel to each other; four parallel veins are present in the specimen. The larger fragment preserves only one margin and six parallel veins.
Remarks. The parallel venation allows a reasonably secure placement within monocots; no more detailed systematic analysis is possible given the fragmentary state of the specimen. Parallel veins without any forking distinguishes it from most cycad or bennettite foliage, which have forked veins.

Palynology
Well-preserved palynological assemblages were recovered from two samples: 47 miospore taxa were recorded, of which 18 taxa constitute spores, 18 gymnosperm pollen and 11 angiosperm pollen. In addition, the freshwater alga Botryococcus occurs in both samples. Most taxa in these samples are long ranging with a mix of typical Jurassic holdovers, such as representatives of Cyathidites, Classopollis, Perinopollenites, Araucariacites and Alisporites, together with typical Late Cretaceous angiosperm pollen. The relatively high abundance of tricolpate and periporate angiosperm pollen (up to 15%) including Normapolles and Alnipollenites sp. (Table 2) suggests an age not older than Campanian and not younger than Maastrichtian, which is consistent with previous age determinations.
Palynofacies. The microscopic organic matter in this sample is dominated (56%) by non-marine palynomorphs and phytoclasts (pollen, spores, wood remains, leaf cuticle and Botryococcus): the rest is represented by dinoflagellates (17%) and AOM (27%: Table 3). No charcoal was identified. The kerogen sample indicates a thermal alteration index (TAI) of 2+, equivalent to 5 on the spore colour index (SCI) chart.

Sample NRM S089755
Sample NRM S089755 consists of fine-grained sandstone to siltstone. It yielded 33 pollen and spore taxa ( Table 2). The spores are represented by 13 taxa and constitute 46% of the assemblage, gymnosperm pollen grains are represented by 15 taxa and attain 37%, and angiosperm pollen (c. 12%) are represented by seven taxa.
As in the previous sample, the spore assemblage is strongly dominated by Cyathidites (31%), followed by Deltoidospora and Gleicheniidites senonicus (Table 2). In this sample, lycophyte spores reach over 4% and are represented mainly by Retitriletes. The relative abundance of the various gymnosperm taxa is more evenly distributed in this sample compared to NRM S089754, and Perinopollenites elatodies dominates with 10%, followed by unidentified bisaccate pollen grains, Classopollis, Alisporites and Araucariacites: all reach relative abundances of about 3%.
Palynofacies. The organic matter in this sample is also dominated by non-marine palynomorphs, but at the significantly higher relative abundance of 80%. Dinoflagellates reach only 4% in this sample, and AOM typical of marine environments reaches only 16% (Table 3). The kerogen sample indicates a TAI of 2+, equivalent to 5 on the SCI chart.

Palaeoecology
The Köpinge flora was unequivocally preserved in a near-shore marine environment, as evidenced by the significant dinoflagellate component in the palynological assemblages and the co-occurrence of abundant marine invertebrates together with the leaf fossils (Fig. 2c). Despite the evidently allochthonous character of the described fossil flora, analogies to better-preserved fossil assemblages allow some general remarks on the Skåne Campanian plant communities. The angiosperm Debeya haldemiana dominates the macroflora in the available collection (Table 1). The apparent dominance of Debeya haldemiana may be biased by the fact that even small fragments of these distinctive leaves may be identified, whereas scraps of other angiosperm leaves cannot. However, even if the true percentage of fossils belonging to this species is perhaps lower, its dominance in the studied assemblage is undeniable. The abundance of Debeya haldemiana in marine sandstones at Köpinge suggests that this species dominated a coastal lowland forest similar to those described from the Cenomanian of Crimea and the Late Cretaceous 'Senonian' of Sakhalin by Krassilov (1979Krassilov ( , 1984; Debeyo-Dryophylletum sensu Krassilov, 1997). Within the palynoflora, however, the gymnosperm pollen grains and the fern spores dominate, which is to be expected as these are wind dispersed and have a much higher spore/pollen production compared to angiosperms, which commonly are insect pollinated. The angiosperm pollen is well represented in the palynoflora, but Debeya pollen is unknown, so no palynomorph can be linked to these megafossils. The fern component is visible in both the spore and the megafloral record.
The local abundance of Perinopollenites elatoides (pollen assigned to Cupressaceae) and Classopollis (pollen attributed to Cheirolepidiaceae) does not show an obvious link to the local megaflora. Although Perinopollenites might be affiliated with the relatively abundant Pagiophyllum, the producers of the Classopollis grains are uncertain. Classopollis derives from cheirolepidiacean conifers, particularly Frenelopsis, which typically occurs in Late Cretaceous marine-influenced habitats (Uličný et al. 1997). This type of conifer is characteristic of Late Cretaceous halophytic habitats (Uličný et al. 1997;Kvaček 2000). However, megafossils of Frenelopsis were not recorded in the Swedish assemblages. This could be due to their lower fossilization potential compared, for example, to Pagiophyllum (cf. Riera et al. 2010). Alternatively, Classopollis might be affiliated with nonfrenelopsid cheirolepids, which could conceivably include some species of Pagiophyllum or similar awl-shaped coniferous foliage (e.g. Otwayia Pole, 2000) based on foliar anatomical characters and associated pollen from both the northern and southern hemispheres (Pole 2000;van der Ham et al. 2003;Tosolini et al. 2015).
By contrast, Ettingshausenia sp., known from a single leaf fragment at the richest locality of Köpinge and from more numerous specimens at Rödmölla and Tosterup (localities closer to the Palaeozoic plateau (Fig. 1c) and, arguably, to the Cretaceous shoreline), is best interpreted as a tree of riparian forests, as generally interpreted for platanoids (Doyle & Hickey 1976;Upchurch et al. 1994;Wing & Boucher 1998;Wing 2000;Coiffard et al. 2006).
It is problematic to link the angiosperm pollen taxa to the megaflora from the Köpinge site and currently we can only speculate. Monocolpate, reticulate Liliacidites pollen (monocotyledonous) is present in the palynoflora, but more abundant are representatives of the Normapolles Group, which make up 12-14% of the miospore assemblages yet are only represented by four taxa. Normapolles is a very diverse group with over 100 described species that had their greatest diversity during the Santonian and Campanian within the northern hemisphere (Skarby 1968;Friis et al. 2006Friis et al. , 2011, but were affected by the Cretaceous -Palaeogene mass extinction event, during which the group lost its diversity (Vajda & Bercovici 2014). The only extant plant-producing Normapolles-like pollen is Rhoiptelea (Friis et al. 2011) within Rhoipteleaceae, Fagales -an order that is represented within the megaflora at Köpinge.

Palaeobiogeography
The Campanian flora of southern Skåne may be compared with three approximately coeval floras: from eastern Poland and western Ukraine (mainly late Campanian: Halamski 2013); from Grünbach in Austria (early Campanian: Kvaček & Herman 2004;Herman & Kvaček 2010); and from Haldem in Westphalia (Hosius & von der Marck 1880), even if only the first two have been revised recently. Debeya haldemiana is a species shared between the floras of Skåne and Poland-Ukraine (although it is not so abundant in the latter regions). Other shared genera (but probably represented by different species) include Ettingshausenia, and probably also Rarytkinia and Korallipteris. Debeya haldemiana is also shared with the Haldem flora (see Halamski & Kvaček 2013), whereas cf. Dryophyllum sp. from Skåne is quite similar to the leaves reported by Hosius & von der Marck (1880) as species of Quercus. It is also notable that Ettingshausenia and probably Korallipteris occur at Haldem (Hosius & von der Marck 1880, pls 38 -39, 42, fig. 176ff ). Distinct similarities between these three floras (Skåne, Poland-Ukraine and Westphalia) reflect analogous palaeoecological settings within the same palaeoclimatic (palaeovegetation) zone. By contrast, the flora described herein is very different from that at Grünbach, which constitutes a thermophilic assemblage from a wet environment.
A comparison between the studied flora and an older (Coniacian) assemblage from the Sudetes (localities Idzików/Kieslingswalde and Nowy Waliszów/Neuwaltersdorf: Kvaček et al. 2015) shows that Korallipteris (earlier described as Microphyllopteris), Debeya, Ettingshausenia, Dryophyllum and Salicites are common to both floras, possibly reflecting similar palaeoecological conditions. Ettingshausenia is characteristic of Late Cretaceous floodplain vegetation (Kvaček & Herman 2004;Greguš & Kvaček 2015). The abundance of conifers in the Köpinge assemblage favours a source from a floodplain swamp vegetation. The occurrence of Classopollis pollen reflects a contribution from halophytic vegetation in the region, although macrofossil elements of this community are not obviously represented in the Skåne megaflora.
Palaeoclimatic inferences based on six dicotyledonous angiosperm taxa can be only very general. However, the approximately equal abundance of entire-margined and serrate/dentate taxa in the studied flora (3:3 if Ettingshausenia sp. is counted as entire-margined and 2:4 if it has a dentate margin) suggests an intermediate character of the vegetation between thermophilic and polar floras. This aspect is consistent with the coeval assemblages from Poland and Ukraine (Halamski 2013, p. 435). The palaeovegetation of a broad transitional zone situated in the European archipelago between approximate palaeolatitudes of 40 -458 N (Poland) and 45-508 N (Skåne) thus seems to have been relatively uniform.
Conclusions † Campanian marine calcareous sandstones from the Vomb Trough in southern Skåne (main locality: Köpinge) yielded a relatively low-diversity vascular flora with one fern, three conifer species (one cone scale and two foliage taxa), six dicotyledonous angiosperms and one monocot angiosperm. † A well-preserved miospore assemblage was recovered from the same sediments as the megaflora, and 47 pollen and spore species were identified. In terms of abundance, the palynoflora is dominated by Cyathidites spores and gymnosperm pollen related to taxodiaceous Cupressaceae. Furthermore, angiosperm pollen is of relatively high diversity and, in terms of abundance, is mainly represented by pollen belonging to the Normapolles complex. † The megafossil assemblage is quantitatively dominated by the angiosperm Debeya (Dewalquea) haldemiana (Deb. ex Sap. & Marion) Halamski [¼D. nilssonii (Brongn. ex Nilsson) Nath. non Sternberg, 1824], which constitutes up to 90% of the specimens. The conifer Pagiophyllum sp. is also a relatively abundant element of the flora. † Neotypes of Dicotylophyllum friesii (Nilsson, 1832) comb. nov. and Salicites wahlbergii (Nilsson, 1832) Hisinger, 1837 are selected. † The palaeoenvironment is interpreted on the basis of the mega-and palynoflora to represent a nearshore, possibly lagoonal, setting. This is more specifically corroborated by the relatively high abundance of dinoflagellates, together with the freshwater to brackish-water alga Botryococcus (which favours calm environments). The angiosperm Debeya haldemiana is interpreted as having formed coastal lowland forests. † The described flora is broadly similar to the approximately coeval assemblages from analogous palaeocommunities described from eastern Poland and western Ukraine (Halamski 2013) and Westphalia (Hosius & von der Marck 1880