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New data on Karksiodus (Chondrichthyes) from the Main Devonian Field (East European Platform)/Uusi andmeid Karksiodus'e (Chondrichthyes) kohta Peadevonivaljalt.


Chondrichthyan remains are very rare in the Devonian shallow-water deposits of the northwestern region of the East European Platform, on the territory called the Main Devonian Field (Borisyak 1922). This territory with the large exposure area of Devonian deposits includes parts of Estonia, Latvia and Lithuania and the northern region of Belarus (the western part of the Main Devonian Field), as well as the northwest of Russia: Leningrad, Pskov, Novgorod and Vologda regions (the eastern part of the Main Devonian Field). Scarce chondrichthyan scales have been reported from the Middle Devonian Narova (= Narva) and Burtnieki regional stages of Estonia and northwestern Russia (Karatajhte-Talimaa 1997; Marss et al. 2008). A few chondrichthyan teeth are also known from the Middle (Arukula and Burtnieki regional stages) and Late Devonian (Kursa and Mhri regional stages) of Latvia, Estonia and northwestern Russia (Ivanov & Luksevics 1994; Mark-Kurik & KaratajhteTalimaa 2004; Ivanov et al. 2011). Karksiodus mirus Ivanov & Marss, 2011 (in Ivanov et al. 2011) is so far the best-known, dentition-based chondrichthyan taxon from the Main Devonian Field. The five teeth ascribed to this species have been recorded in the Givetian Arukula deposits of the Arukula cave (Mark-Kurik & Karatajhte-Talimaa 2004) and Burtnieki deposits of the Karksi outcrop, Estonia (Ivanov et al. 2011). The additional teeth from the Karksi outcrop and the new specimens from the Givetian of the localities from the Leningrad Region, Russia, are described in this paper.


The teeth of Karksiodus collected to date are usually incomplete, with broken cusps and bases, exposing the walls of the transverse basal vascular canal; they also often show abraded surfaces. However, some teeth bear well-preserved striations on the cusps.

Most of the sandstone layers containing chondrichthyan remains were very weakly cemented and did not require acid preparation. Samples of the sandstone were fractioned and sieved. Vertebrate microremains were then manually picked under an optical microscope. Acetic acid was utilized to recover vertebrate microremains from the samples of dense, carbonate-cemented sandstone from the Kemka locality. The tooth microstructure was examined in thin section; the images were photographed at the different stages of polishing using the optical microscope Nikon ECLIPSE 50i and digital camera Nikon DS-Fi1. One tooth cusp was polished and etched for 10 sec in 5% HCl. The microremains were micrographed using the scanning electron microscopes Cambridge CamScan-4, Tescan VEGA-II XMU and Zeiss EVO MA15.

The described specimens are housed at the Paleontological Museum of St. Petersburg State University (collection number PM SPU 70) and in the Institute of Geology at Tallinn University of Technology (collection number GIT 383 with specimen numbers 47-49).


The new teeth of Karksiodus mirus Ivanov & Marss were found in five distinct localities within the Leningrad Region, northwestern Russia (Fig. 1): Lemovzha (Volosovo District), Siverskij, Zaitsevo and Novinka (Gatchina District) and Kemka (Luga District).

Lemovzha locality

The outcrop is located on the right bank of the Lemovzha River (right tributary of the Luga River), 4.5 km upstream from the river mouth and 2.5 km from Khotnezha village (Fig. 1B). The Middle Devonian beds are well exposed in a cliff about 20 m high and 40 m long. The section includes the deposits of the Narova (Narva) Regional Stage in the lower part of the outcrop and the Arukula Regional Stage (lower part of the Luga Regional Beds) in the upper part. The Narova deposits of the Lemovzha and Khotnezha members are represented by brownish or dappled marls with brownishgrey sandstones, bluish-grey siltstone and clay interlayers. Some marl levels downstream along the Lemovzha River contain well-preserved remains of agnathans and fishes (Ivanov & Lebedev 2011; Skutschas et al. 2011; Ivanov et al. 2012). The vertebrate assemblage occurring there includes the psammosteids Schizosteus splendens (Eichwald) and S. striatus (Gross); the placoderms Holonema sp., Homostius sp. and Byssacanthus dilatatus (Eichwald); the acanthodians Archaeacanthus sp., Cheiracanthus longicostatus Gross, C. brevicostatus Gross, Diplacanthus sp., Haplacanthus sp., Markacanthus costulatus Valiukevicius, Ptychodictyon sp. and P. distinctum Valiukevicius; and the sarcopterygians Glyptolepis? quadrata Eichwald, Struniiformes indet., Dipterus arenaceus Eichwald, cf. Gyroptychius sp., Thursius sp. and Osteolepiformes indet. (Valiukevicius 1985; Ivanov et al. 2012, with additional data herein).

The deposits of the Arukula Regional Stage are represented by brownish- and yellowish-grey, fine- to coarse-grained, cross-bedded sandstones with clay balls, lenses and interlayers. The sandstones are moderately cemented with weakly cemented interlayers. The Arukula vertebrate assemblage known from the surroundings of Khotnezha village contains remains of the psammosteids Pycnosteus palaeformis Preobrazhensky, P. pauli Mark and Schizosteus striatus; the placoderms Homostius sp., Actinolepis tuberculata Agassiz, Dickosteus sp. and Asterolepis estonica Gross; the acanthodians Archaeacanthus quadrisulcatus Kade, Haplacanthus marginalis Agassiz, Homacanthus gracilis (Eichwald), Diplacanthus carinatus Gross, D. gravis Valiukevicius, Diplacanthus sp., Markacanthus costulatus Valiukevicius, M. alius Valiukevicius, Minioracanthus laevis Valiukevicius, Cheiracanthus brevicostatus Gross, C. longicostatus Gross, C. talimae Valiukevicius and Ptychodictyon sp.; and the sarcopterygians Glyptolepis sp., Dipterus radiatus (Eichwald) and Osteolepididae indet. (Valiukevicius 1985; Ivanov & Lebedev 2011, with additional data herein). The psammosteids and sarcopterygians are predominant among the vertebrate macroremains.

The precise position of the boundary between the Narova and Arukula regional stages has not been determined in the Lemovzha section. The tooth of Karksiodus mirus Ivanov & Marss was found in the layer of brownish-grey, weakly cemented, fine- to medium-grained, cross-bedded sandstones, 1.5 m above the marls and sandstones boundary. This layer contains remains of the psammosteids Schizosteus splendens, S. striatus and Pycnosteus pauli (V. Glinskiy, pers. comm. 2013); the placoderms Asterolepis sp. and Coccosteidae indet.; the acanthodians Cheiracanthus brevicostatus, Diplacanthus gravis, 'Acanthodes' sp., Homacanthus sp. and Rhadinacanthus sp.; and the sarcopterygians Glyptolepis sp. and Osteolepididae indet. The psammosteid taxa S. striatus and P. pauli occur in the Narova and Arukula regional stages. However, S. splendens is only found within the Narova Regional Stage (Mark-Kurik 2000). On the other hand, the acanthodian species Diplacanthus gravis is only known to occur in the Arukula Regional Stage (Mark-Kurik 2000; Valiukevicius 2000). For this reason, it is likely that the lower sandstone layer containing the Karksiodus tooth is of Arukula age. Numerous and diverse acanthodian scales are also concentrated within this layer, as well as scales of Karksilepis parva Marss. The scale surface has four elongated odontodes that point sharply towards the posterior margin of the scale and a rhomboid, flat base (Fig. 2L). The slightly flattened, horizontally directed odontodes have a smooth surface, but their basal part bears fine striations (Fig. 2L, sc), as typical of Karksilepis (Marss et al. 2008). The central and largest of the odontodes extends across the entire base of the scale and projects beyond the posterior edge of the scale. Two small lateral odontodes are distributed in a row on either side of the central odontode and partly overlap.

Apart from the Karksi outcrop, the Lemovzha outcrop is the second locality where the teeth of Karksiodus and scale of Karksilepis occur jointly.

Siverskij locality

The locality is situated on the right bank of the Oredezh River, in Siverskij village, near Siverskaya railway station, at the dam of a former hydroelectric power station (Fig. 1B). The outcrop is about 10 m high and 60 m long, one of a series of outcrops in the valley of the Oredezh River from Siverskaya to Belogorka villages (Ivanov et al. 2005). The deposits of the outcrops belong to the lower part of the Arukula Regional Stage and are represented by striped yellow, brown and orange, fine- to medium-grained, poorly cemented to dense, micaceous-quartz sandstones with conglomerate intercalation. The sandstones contain clay balls, up to 8 cm in diameter, which form the interlayer in the upper part of the outcrop. Cross-stratifications dominate the section, while planar and wedge-shaped stratifications are rare. Three long lentiform conglomerate interlayers are exposed in the middle part of the section. They are variegated, with dominant yellowish and brownish colours, and contain siltstone balls and silty clay, quartz gravel and coarse sand. The interlayer boundaries are wavy and sharp. These conglomerate interlayers, as well as some levels of sandstones, contain the vertebrate remains in different states of preservation.

The vertebrates are mainly represented by isolated skeletal elements and their fragments, but well-preserved tiny remains are also common. The almost complete skeleton of the placoderm Heterostius ingens Asmuss is found at the same level in the Arukula deposits near Belogorka village (Ivanov et al. 2005). The vertebrate assemblage of this locality belongs to the Pycnosteus palaeformis psammosteid Zone. It includes the psammosteid agnathans Pycnosteus palaeformis Preobrazhensky, Schizosteus cf. S. asatkini Obruchev, S. striatus (Gross), Tartuosteus giganteus (Gross) and Psammolepis proia Mark-Kurik; the placoderms Actinolepis tuberculata Agassiz, Heterostius ingens Asmuss, Homostius latus Asmuss, Coccosteidae, Asterolepis sp. and Byssacanthus sp.; the acanthodians Haplacanthus sp., Cheiracanthus cf. C. brevicostatus Gross, Diplacanthus sp., 'Acanthodes' sp. and Homacanthus sp.; and the sarcopterygians Glyptolepis sp., Dipterus radiatus (Eichwald), Gyroptychius pauli Vorobyeva and Thursius sp. (Ivanov et al. 2005, 2012; Glinskiy 2012). Psammosteids, placoderms and sarcopterygians dominate in this vertebrate assemblage. The teeth of Karksiodus mirus Ivanov & Marss were found at two levels: one tooth in the sandstone layer between two conglomerate interlayers, the other in the sandstone layer above the upper conglomerate interlayer.

Zaitsevo locality

The Zaitsevo quarry is located 700 m east of Zaitsevo village, 1.2 km east of Stroganovo railway station, Gatchina District (Fig. 1B). The deposits of the Arukula Regional Stage are represented here by brownish, pinkgrey and yellow-grey, fine- to medium-grained, commonly weakly cemented to rarely dense, micaceous-quartz sandstones with interlayers and a lens of yellowish-grey clays and siltstones, with silty clay balls, up to 12 cm in diameter. Cross-stratifications are dominant through the section, while planar stratifications with concentration of oriented clay balls are rare. The densely cemented sandstones in the upper part of the section have a thin laminar structure and contain trace fossils.

The vertebrate remains are commonly represented by isolated skeletal elements and their fragments. The assemblage comprises the psammosteids Pycnosteus palaeformis Preobrazhensky, Tartuosteus giganteus (Gross) and Psammolepis proia Mark-Kurik (Glinskiy 2012); the placoderms Actinolepis tuberculata Agassiz, Heterostius ingens Asmuss, Homostius latus Asmuss, Coccosteidae, Asterolepis sp. and Byssacanthus sp.; the acanthodians Haplacanthus sp. and Homacanthus sp.; and the sarcopterygians Glyptolepis sp., Holoptychiidae and Gyroptychius pauli Vorobyeva. The psammosteids and placoderms dominate in this vertebrate assemblage of the Pycnosteus palaeformis psammosteid Zone of the Arukula Regional Stage. The tooth of Karksiodus mirus Ivanov & Marss was found in brownish-grey, weakly cemented sandstone with clay balls.

Kemka locality

A series of Middle Devonian outcrops are exposed on both banks of the canyon-like valley of the Kemka River (right tributary of the Luga River), 9 km upstream from the river mouth, in the middle course and between the mouths of Domanov and Lobovoj creeks (Fig. 1B). The tooth of Karksiodus mirus Ivanov & Marss was collected in one of these outcrops, on the right bank of the river. The outcrop is about 7 m high and 40 m long. The deposits belonging to the lower part of the Burtnieki Regional Stage comprise pinky-yellow or brownish, fine- to coarse-grained, cross-bedded sandstones containing clay balls with interlayers of bluish-grey or yellow siltstones, mudstones and clays. The sandstones are normally weakly cemented but some dense interlayers with carbonate cement are also found.

The vertebrates are represented by their isolated skeletal elements and fragments but parts of the skeleton are rarely recorded from there. The ichthyoassemblage belongs to the Pycnosteus tuberculatus psammosteid Zone and the Asterolepis delli placoderm Zone, and includes the psammosteids Pycnosteus tuberculatus (Rohon), Ganosteus stellatus Rohon (Glinskiy 2012); the placoderms Actinolepis sp., Homostius latus Asmuss, Holonema sp., Dickosteus sp., Asterolepis cf. A. delli Gross; the acanthodians Haplacanthus sp., Ptychodictyon sp., 'Acanthodes' sp.; the sarcopterygians Glyptolepis sp., Porolepiformes, Dipteridae, Osteolepididae; and the actinopterygian Cheirolepis sp. The placoderm remains are abundant in the assemblage. The Karksiodus tooth was found within a carbonate-cemented, dense sandstone interlayer.

Novinka locality

The Novinka quarry is located 1.1 km south of Novinka village (railway station), Gatchina District (Fig. 1B). The deposits of the Burtnieki Regional Stage include yellow-grey, fine- to medium-grained, commonly weakly cemented, micaceous-quartz sandstones with interlayers of yellowish-grey and bluish-grey clays and siltstones (Ivanov et al. 2012).

Vertebrate remains have been collected from pinkishgrey and red medium-grained trough-cross-stratified sandstones. The vertebrates are represented mainly by isolated skeletal elements and fragments of bones, plates, fin spines, scales and teeth. The diverse agnathan and fish remains include the psammosteids Pycnosteus tuberculatus (Rohon), Pycnosteus sp., Ganosteus stellatus Rohon, Tartuosteus maximus Mark-Kurik, Psammolepis abavica Mark-Kurik and Psammosteus bergi (Obruchev); the placoderms Rhynchodus sp., Actinolepis magna Mark-Kurik, Homostius cf. latus Asmuss, Heterostius sp., Dickosteus sp. and Asterolepis delli Gross; the acanthodians Homacanthus sp., 'Acanthodes' sp. and Nostolepis sp.; and the sarcopterygians Glyptolepis sp., Porolepiformes indet., Conchodus sp., Dipnoi indet. and Osteolepididae indet. (Ivanov & Glinskiy 2011). Psammosteid and large arthrodiran remains are dominant in this assemblage. This assemblage belongs to the Pycnosteus tuberculatus and Asterolepis delli zones. The teeth of Karksiodus mirus Ivanov & Marss were found in the upper layer of yellowish-grey siltstones.

Karksi locality

Apart from the four teeth of Karksiodus mirus Ivanov & Marss described earlier (Ivanov et al. 2011), three additional incomplete teeth have been found in the Harma Beds, Burtnieki Regional Stage of the Karksi outcrop in South Estonia (Fig. 1A). The teeth were collected in the samples from two levels, 02-1 and 90 (= 02-4) of this outcrop (Marss et al. 2008). Besides the teeth of the elasmobranch Karksiodus and the scales of the putative chondrichthyan Karksilepis parva Marss, the vertebrate assemblage from the Karksi locality also contains the scales of typical chondrichthyans. One scale from sample 02-4 (Fig. 3C, D) possesses a rhomboid crown with a slightly acuminated posterior part. The crown is shallowly inclined anteriorly and projected over the scale base. The anterior part of the external crown surface bears 11 narrow and long ridges. These are separated by deep depressions anteriorly, are smooth in the middle part and do not reach the smooth crown posteriorly. The ridges are of different length and some of them are branched posteriorly. The anterior edge of the crown is uneven and sinuous. The scale neck is not well developed anteriorly but is deeply constricted posteriorly. The base is low and smaller than the crown, with its upper surface rising towards the crown and having a flat basal surface. The second scale, from sample 97 (Fig. 3E) is incompletely preserved. It has a small, slightly inclined crown, narrow neck and large oval base. The crown surface is flat, with an uneven anterior edge. The base has a pyramidal upper surface and a slightly concave basal surface.


External morphology

Compared with the already described specimens, the new collection of Karksiodus teeth provides additional and systematically valuable characters, such as a variable dental morphology. The range of tooth sizes is substantial: the width of the base varies from 0.5 mm (Fig. 2C, D) to 2.3 mm (Fig. 2J, K), mostly measuring 1.2-1.6 mm.

The tooth crown includes two to four cusps. The bicuspid tooth (Mark-Kurik & Karatajhte-Talimaa 2004, fig. 3A) bears two large lateral cusps on the crown. The more common tricuspid teeth possess two main lateral cusps with a small central cusp displaced more labially than the lateral ones. The crowns of the four-cusped teeth have an additional intermediate cusp (cusplet) placed either between the central and lateral ones or in line with the central cusp (Fig. 2J, K), or placed more medially in line with both lateral cusps (Fig. 3A, B). The intermediate cusp, as observed in one broken tooth, is either narrower than the central cusp or of the same width. Cusp striations among the known teeth of Karksiodus vary from very fine and dense to coarse and sparse. The cusps are usually rounded in cross section but sometimes oval and flattened labio-lingually. The angles between the lateral cusps range from 40[degrees] to 60[degrees], the largest angle occurring in the bicuspid and multicuspid teeth; the lateral cusps of tricuspid crowns diverge at an angle of about 50[degrees].

The structure of the base of all known teeth of Karksiodus varies largely. The tooth base shows various degrees of curvature, from slightly curve (Fig. 2A-D, J, K; Mark-Kurik & Karatajute-Talimaa 2004, fig. 3A) to strongly arcuated (Fig. 2E, G; Ivanov et al. 2011, fig. 3A-C). The bicuspid, multicuspid and some small tricuspid teeth have the flattest bases. The length of the lateral prominent parts (projections) of the tooth base can vary from short (Fig. 2K) to considerably long (Fig. 2E). These projections are well developed in the large teeth with strongly arched bases. The prominence of the transverse basal vascular canal wall on the basal surface of the tooth varies from not being salient to strongly salient. In the former case the external wall of the canal is contained within the smooth basal surface. The tricuspid teeth with strongly arched bases and extended lateral projections possess the most prominent transverse basal canal wall. In the bicuspid, multicuspid (Fig. 2J, K) and small tricuspid teeth (Fig. 2C, D) this canal is only slightly protruded. The transverse basal canals commonly run along the entire midline of the base, but they can have an asymmetrical position, disposed from the midline either mesially or distally (Fig. 2G; Ivanov et al. 2011, fig. 3F, I).

Internal structure

A longitudinal thin section of a tooth (Fig. 4 A) and fragmented teeth from the new collection add some new features of the internal structure of the Karksiodus teeth. Orthodentine tubules make up the lateral cusps from the base to the cusp apex (Fig. 4D), but they are more densely concentrated and wider at the cusp/base boundary (Fig. 4C). The boundary between the orthodentine of the cusps and the trabecular dentine of the base is distinct. Some of the cusps of the new specimens have well-preserved external surfaces but no enameloid layer was detected in any of the teeth, even during the SEM study of the cross section of the tooth cusp polished and acid-etched. Probably the external layer of the cusp forming the striation consists of hypermineralized superficial orthodentine.

The vascularization system of the Karksiodus teeth includes four types of vascular canals: transverse basal canals (Fig. 4B, tbc), thin and small canals forming the network within the base (ascending and horizontal, Fig. 4B, avc and hvc), rare large canals (Fig. 4B, lvc) and pulp canals within the lateral cusps (Fig. 4D, pc). The external wall of the tube-like transverse canal consists of compact tissue as mentioned by Ivanov et al. (2011) and is not perforated by small canals. The inner surface is penetrated by several foramina of horizontal and ascending small vascular canals connecting the transverse canal with its network. One large vascular canal opens on the lingual rim of the tooth base, alongside the foramen of the transversal canal (Figs 2E, G; 3B, lvc). The large canal is subdivided within the base into two branches and runs sub-parallel with the basal surface (Fig. 4B, lvc). This canal is connected with thin canals. The thin ascending vascular canals are graded to the broadened pulp canal of the lateral cusp (Fig. 2I). The teeth of Karksiodus exhibit a compound vascularization system.


To summarize the already known and new data, Karksiodus mirus Ivanov & Marss occurs in the Givetian, Middle Devonian of South Estonia: the Arukula Regional Stage of the Arukula cave, Tartu (Mark-Kurik & Karatajhte-Talimaa 2004), and the Burtnieki Regional Stage of the Karksi outcrop (Ivanov et al. 2011), and in the Leningrad Region, Northwest Russia: the Arukula Regional Stage of the Lemovzha, Siverskij and Zaitsevo localities, and the Burtnieki Regional Stage of the Kemka and Novinka localities.

In Karksiodus, variations in tooth morphology, such as the curvature of the base, length of lateral parts, prominence of the transverse basal canal and details of cusp external sculpture (striations) are possibly related to different stages of tooth growth as observed in the tricuspid teeth of various sizes. However, the variability in the crown structure (number of cusps and angles between the lateral cusps), as well as some variability in the base (position of the basal canal and possibly base curvature), suggest that Karksiodus dentition displayed weakly developed heterodonty in which the bicuspid, tricuspid and multicuspid teeth were located in different positions on the jaws.

With the new tooth material, the arrangement of the tooth rows can be tentatively reconstructed. The teeth would not extensively overlap each other but rather the lingual rim of each tooth would be in contact with the shallow longitudinal groove on the labial side of the tooth in front; the basal surface of adjacent teeth in the same row would abut on each other forming an archlike structure.

Nevertheless, the new material has not provided enough evidence to clarify the interrelationships of Karksiodus with other chondrichthyans. The variations in the crown structure, especially the changeable number of intermediate (including central) cusplets, resemble the tooth structure of the Doliodus and the Antarctilamna-Wellerodus group. The latter differs from Karksiodus in having a lingual extension of the base and elements for tooth-to-tooth articulation, including the apical button and labio-basal projection (Ginter et al. 2010). On the other hand, Doliodus dentition includes commonly the tooth row with fused bases but the isolated teeth possess labio-basal extensions of the base (Turner 2004; Maisey et al. 2014). None of these groups display the same histological characters presented by Karksiodus, namely, the complex vascularization system made up of four distinct canal types, which is so far unique to Karksiodus among either extinct or extant chondrichthyans.

doi: 10.3176/earth.2014.14

Acknowledgements. The authors are grateful to Vadim Glinskiy (St. Petersburg State University), Anne Kleesment (Institute of Geology at Tallinn University of Technology (TUT)) and Alexey Kuzmin (Karpinsky Russian Geological Research Institute, St. Petersburg) for kindly providing microremain materials for this study, to Vadim Glinskiy for unpublished identifications of psammosteid taxa, to Pavel Skutschas (St. Petersburg State University) for data on the Lemovzha locality, to Rutt Hints (Institute of Geology at TUT) and Roman Rakitov (Borissiak Palaeontological Institute of the Russian Academy of Sciences, Moscow) for assistance during SEM imaging and to Gennadi Baranov (Institute of Geology at TUT) for improving some illustrations. We thank Mags Duncan (Ireland) for improving our English and Martha Richter (UK) for review and text correction. T.M. was supported by the Estonian Target Funding Project SF0140020s08. A.I. acknowledges St. Petersburg State University for research grant This work was partially funded by the subsidy of the Russian Government to support the Program of Competitive Growth of Kazan Federal University among World's Leading Academic Centres. The study was supported by RFBR, research project No. 14-04-01507a.


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Alexander Ivanov (a,b) and Tiiu Marss (c)

(a) Faculty of Geology, St. Petersburg State University, 16 Liniya 29, 199178, St. Petersburg, Russia; IvanovA-

(b) Institute of Geology and Petroleum Technology, Kazan Federal University, Kremlevskaya St. 4/5, Kazan, Russia

(c) Institute of Geology at Tallinn University of Technology, Ehitajate tee 5, 19086 Tallinn, Estonia;

Received 25 May 2014, accepted 13 August 2014
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Author:Ivanov, Alexander; Marss, Tiiu
Publication:Estonian Journal of Earth Sciences
Article Type:Report
Geographic Code:4EXES
Date:Sep 1, 2014
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