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Karksilepis parva gen. et sp. nov. (Chondrichthyes) from the Burtnieki Regional Stage, Middle Devonian of Estonia/Karksilepis parva gen. et sp. nov. (Chondrichthyes) Kesk-Devoni Burtnieki lademest Eestis.

INTRODUCTION

Chondrichthyan scales are rare in Lower-Middle Palaeozoic rocks. Chondrichthyan-like scales, however, were described by Sansom et al. (1996) from the Ordovician of Colorado, USA, and by Young (1997) from central Australia. The earliest Elasmobranchii gen. nov. has been listed from the Early Llandovery (Early Silurian) of the Siberian Platform, Russia (Karatajute-Talimaa & Predtechensk)j 1995). Niualepis and Elegestolepis conica have been identified in the Middle Llandovery of South Yakutia and the Bratsk Region, Siberian Platform (Novitskaya & Karatajute-Talimaa 1986; Karatajute-Talimaa 1992). Elegestolepis sp. comes from the Upper Llandovery of Irkutsk Amphitheatre, Tuva, and North Mongolia (Vladimirskaya et al. 1986).

Scales of an elasmobranch (Kannathalepis) and possible chondrichthyans (Frigordepis and Wellingtonella) originate from the Wenlock of the Canadian Arctic (Marss & Gagnier 2001; Marss et al. 2002, 2006). Elegestolepis grossi has been recognized in the Upper Silurian (Upper Ludlow-Pridoli) of Tuva (Karatajute-Talimaa 1973). Four chondrichthyan elasmobranch(?) genera, Seretolepis, Altholepis, Knerialepis, and Ivanelepis, established on the basis of scales, have been described from the Lochkovian of Podolia (Obruchev & Karatajute-Talimaa 1967; Karatajute-Talimaa 1968, 1977, 1997a; Hanke & Karatajute-Talimaa 2002). Polymerolepis Karatajute-Talimaa was first discovered in the Lochkovian of Podolia (Obruchev & Karatajute-Talimaa 1967). Later on it has also been identified in the coeval strata of Central Nevada (Turner & Murphy 1988) and on Ellesmere and Prince of Wales islands of the Canadian Arctic (Langenstrassen & Schultze 1996). The list of chondrichthyan and putative chondrichthyan taxa from the Lochkovian of the Mackenzie Mountains (MOTH locality), Canada, includes Kathemacanthus, Altholepis, Seretolepis, Polymerolepis species, and five unnamed taxa (Hanke & Wilson 1997, 1998; Wilson & Hanke 1998; Wilson et al. 2000). Ellesmereia are found in the Lochkovian of the Canadian Arctic (Vieth 1980). Scales of Arauzia, Lunalepis, and Iberolepis are known from the Lower Devonian, Lower Gedinnian (Lochkovian) of Spain (Mader 1986). Pamyrolepis nom. nud. Karatajute-Talimaa scales have been briefly described from the Emsian of the Eastern Pamyrs (Karatajute-Talimaa 1992; Leleshus et al. 2005).

Mongolepidids were first treated as belonging to Elasmobranchii, Chondrichthyes (e.g. Karatajute-Talimaa et al. 1990). Later on, Karatajute-Talimaa (1995) suggested that Mongolepidida is an independent group of lower vertebrates, still related to the Chondrichthyes. The group is rather widely known in the Silurian. Mongolepis, Sodolepis, and Teslepis have been described in the Upper Llandovery of West Mongolia (Karatajute-Talimaa et al. 1990; Karatajute-Talimaa 1992; Karatajute-Talimaa & Novitskaya 1992, 1997). Several taxa belonging to the mongolepidids come from the Upper Llandovery of different sites of China: Xinjiangichthys from Kalpin and Bachu counties, Xinjiang, Shiqianolepis and two taxa of uncertain affinity, Rongolepis and Chenolepis, from Shiqian county, Guizhou Province (Wang et al. 1998; Sansom et al. 2000). In the Wenlock (Lower Silurian) of the Siberian Platform Mongolepididae gen. nov. B has been identified together with Elegestolepis? sp. (Karatajute-Talimaa & Predtechenskyj 1995). The latest mongolepidids of the world were discovered in the probable Lochkovian (Lower Devonian) strata of Eastern Mongolia (Wrona & Nyamsuren 1998) and in the Lower Emsian (Lower Devonian) of Gornyi Altai of Western Siberia, showing the wide stratigraphical range of the whole group (Rodina 2002).

Scales of chondrichthyans are rare in the East Baltic. Lugalepis multispinata comes from the Emsian(?) of Latvia (Karatajute-Talimaa 1997b), while the Eifelian (Middle Devonian) Narva Regional Stage (for short, RS) of Leningrad District, Russia, Belarus, and Lithuania contains a somewhat different form Lugalepis cf. multispinata (Karatajute-Talimaa 1992, 1997b). Rare chondrichthyan buccopharyngeal scales (= oral denticles) and teeth have been documented from the Givetian (Middle Devonian) of Estonia and from the Famennian (Upper Devonian) of Latvia (Ivanov & Luksevics 1994; Mark-Kurik & Karatajute-Talimaa 2004).

In the course of lithological studies of Devonian sediments in Estonia, based upon outcrops and drill core material, a rich collection of vertebrate microfossils has been obtained. Notably abundant material has been collected from the Karksi outcrop in the Halliste old valley, near Karksi castle (Viljandi county) (Fig. 1). The sandstones here belong to the lower part of the Harma Beds of the Burtnieki RS and are famous for fish fossils. Vertebrates described in earlier studies include psammosteids (Obruchev & Mark-Kurik 1965; Mark-Kurik 1968, 1995), placoderms (Karatajute-Talimaa 1963), acanthodians (Valiukevicius 1998), and sarcopterygians (Vorobyeva 1977). Chondrichthyans have not been mentioned or described so far from these beds.

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In this paper we describe scales of a new chondrichthyan from the Karksi outcrop, Estonia. The authorship of the new taxon Karksilepis parva is attributed herein to Marss.

MATERIAL AND METHODS

Six samples (200-400 g) were collected from four levels of the Karksi outcrop (Fig. 2) and examined. All samples, those previously collected by A. Kleesment in 1990 and 1997 and the ones from two new levels (samples 02-1 and 02-2) contained abundant vertebrate microremains. Scales and tesserae or fragments of plates of heterostracans, placoderms, acanthodians, chondrichthyans, and osteichthyans (sarcopterygians and actinopterygians) were found (see Table 1). Psammosteids predominated, especially plate fragments and scales of Tartuosteus maximus, Pycnosteus tuberculatus, and Ganosteus stellatus. The diversity of acanthodians was high (16 taxa), and Nostolepis gaujensis scales were found for the first time in the Burtnieki RS (they were absent only in sample 02-3). Chondrichthyans were fairly rare but present in all six samples. In addition to the scales of the new taxon described herein, the samples contained teeth of an elasmobranch. As there was no indication that these teeth might belong to the taxon described below (except their co-occurrence in the samples from the same outcrop), they will be described elsewhere. Scales, plate fragments, and teeth of Sarcopterygii were found as well. Actinopterygians were represented by rare scales of Cheirolepis sp. (Kleesment et al. 2003; Marss et al. 2003).

The sandstones in the Karksi outcrop were very weakly cemented and did not need acid preparation. Samples were carefully washed to remove muddy (<0.01 mm) particles. Then the material was fractioned and sieved; the size ranges were 0.1-0.25 mm, 0.25-0.5 mm, 0.5-1 mm, 1-2 mm, and >2 mm. The two finest fractions were separated to heavy and light parts using bromoform (2.89 g/[cm.sup.3]). The two heavy fractions obtained, and the two coarsest fractions were investigated using the binocular optical microscope. All microremains with well-preserved sculpture and shape were picked out. The microstructure of chondrichthyan scales was examined in thin sections or by submerging elements into immersion liquids. A selection of scales was photographed using the scanning electron microscope JEOL-JSM-940-A. The described collection is housed in the Institute of Geology at Tallinn University of Technology under the collection numbers GIT 383 + specimen number.

GEOLOGICAL BACKGROUND

The Middle Devonian Burtnieki RS is distributed in southeastern Estonia (Fig. 1). The total thickness of the stage is 60-95 m. The sequence is represented by light (white, yellowish, pinkish, and greyish-brown) fine-grained weakly cemented cross-bedded sandstones with siltstone and clay interlayers. The stage is divided into three successive cyclic sandstone complexes, the Harma, Koorkula, and Abava beds (Kleesment 1995; Kleesment & Mark-Kurik 1997). The lower unit, the Harma Beds, is 13-28 m thick. The beds cropping out in Karksi belong to the lower part of the Harma Beds. The cross-bedded sandstones of the Harma Beds with the stratotype section in Harma settlement at the Ohne River are usually well sorted and fine-grained, where the size fraction 0.1-0.25 mm clearly dominates (75-93%; Kleesment 1995). The sandstones in the Karksi outcrop and coeval sediments at the Ahja River and in the Torva area have the same composition as the Harma Beds. Sandstones of the Harma Beds possibly accumulated in regressive conditions of the Devonian basin in a fluvially dominated subaqueous delta plain environment, which was under influences of low-amplitude eustatic and tectonic movements, repeatedly interrupted by short subaerial periods (Kleesment 1997; Plink-Bjorklund & Bjorklund 1999).

The part of the Karksi outcrop that yielded fish microremains is a small sandstone scarp 3.5 m high and 6 m long. About 20 m upstream, in the upper part of the slope there is another, smaller fish-bearing locality. The levels with abundant fish microremains are represented by moderately sorted fine- to medium-grained sandstones which contain both particle types in equal amounts and up to 10% coarse and very coarse detrital material. In the coarse-grained (diameter 0.5-2 mm) detrital material, heterostracan, placoderm, and sarcopterygian plate fragments are fairly worn. Acanthodian and chondrichthyan scales are concentrated in beds with the grain size of 0.2-0.5 mm and are not worn, which gives evidence of more rapid accumulation of sediment. Probably the reworking and weathering of fish bones and plates took place in shore face environments, where under low-amplitude eustatic movements, sea level fluctuations caused repeated alternation of marine and non-marine conditions and intense redeposition and reworking of the detrital component. Interlayers with rich, well-preserved fish microremains possibly deposited in the nearshore realm during short periods of intense influx of detrital material. Such rapid burial in nearshore deltaic conditions resulted in the accumulation of well-preserved fish microremains. In these layers, besides moderate sorting of detrital material among fine-grained particles, subrounded to subangular quartz grains dominate (85-90%); in the coarse-grained fraction, rounded grains make up only 25-30%, referring to intense influx and rapid deposition. Such high concentrations of fish remains have been reported from nearshore marine (Davies et al. 2007), deltaic (Mancuso 2003), and lake deposits (Smith & Swart 2002).

SYSTEMATIC PALAEONTOLOGY

Class Chondrichthyes

Order et Family incertae sedis

Karksilepis gen. nov. Marss

Derivation of name. From Karksi locality yielding scales of this taxon, and lepis, meaning scale, gender feminine.

Type species. Karksilepis parva gen. et sp. nov. Marss

Diagnosis. As for type species.

Karksilepis parva gen. et sp. nov. Marss

Figures 3-5

Chondrichthyes gen. et sp.: Marss 2006, figs 7A, B.

Derivation of name. Latin parvus, meaning small, an allusion to the size of the scales.

Holotype. GIT 383-4, scale in Fig. 3B, and its details in Fig. 3C, D.

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Type locality and horizon. Karksi outcrop (sample 97), South Estonia; Harma Beds, Burtnieki Regional Stage, Middle Devonian.

Material. 50 scales, fairly well preserved.

Occurrence. In four levels in Karksi outcrop (see Table 1 and Fig. 2).

Diagnosis. Scales very small to medium-sized, quadrangular (rectangular to rhomboidal), 0.25-0.8 mm long and 0.25-0.7 mm wide. Three to eighteen elongate, partly overlapping longitudinal odontodes on the scale surface; odontode width 0.05-0.1 mm and length 0.15-0.4 mm. Lower face of basal plate convex. Very fine dentine tubules in odontodes branch from wider dentine canals, and the latter in turn from pulp canal; vascular canals form network beneath odontodes in the basal plate; openings of vascular canals situated around odontodes. Flattened bone cell lacunae form lamellar structure of the basal plate. Sharpey fibres very fine.

Description

Morphology. The scales are small to medium-sized, quadrangular, with the shape varying from high rectangle to relatively low rhomboid (Fig. 3A, B, E-K, N, P, Q). The length of the scales is 0.25-0.8 mm and width 0.25-0.7 mm. The height of the basal plate is about half the height of the scale (Fig. 3L). The scales are covered with odontodes forming polyodontodia. The number of odontodes varies from three (Fig. 3Q) to eighteen (Fig. 3J); their width is 0.05-0.1 mm and length 0.15-0.4 mm. The odontodes are attached to the basal plate of the scale. The rectangular high scales carry odontodes in one to two rows (Fig. 3A, B, E, G); the odontodes on rhomboidal scales are less regularly arranged in rows but still up to four rows can be counted (Fig. 3H, J, N), whereas more anterior odontodes partly cover the posterior ones. There is always lateral separation between adjacent odontodes (e.g. Fig. 3A, E, K). The odontodes can be directed horizontally (Fig. 3M) or rise to the posterior (Fig. 3K; see also Fig. 4). All odontodes are covered with fine ultrasculpture of longitudinal striation which starts at the conjunction of the odontode and basal plate and is directed upwards and posteriorwards, towards the peak of the odontode (Fig. 3A-E, G, I, K, M). The basal plate is larger than the area with odontodes. In some scales the basal plate strongly protrudes anteriorly (Fig. 3A). Relatively large deep pores open in the basal plate (Fig. 3G, K-M, P, Q) and make the surface of the plate uneven. The shape of the basal plate depends on the age of the scale. Vascular network canals open anteriorly of the odontodes in high rectangular scales (Fig. 313, E, G) or around the odontodes in rhomboidal scales (Fig. 3K-M, P, Q). The lower surface of the basal plate can be either concave, smooth (Fig. 3F), or convex (Fig. 3L, M, O).

A few rhomboidal to roundish scales have their basal plates covered with several (two to seven) odontodes with a flat and smooth surface. The length of these scales is 0.45-0.75 mm and width 0.3-0.7 mm (Figs 3S-U; 5D). The width of the odontodes is 0.1-0.2 mm and length 0.2-0.4 mm. The odontodes are oval (Fig. 3T, U) to oblong (Fig. 3S), more than twice as long as their width. The odontodes lie close to each other (Fig. 3T, U), some space may be between the anterior odontodes and the neighbouring ones (Fig. 3S). Anterior odontodes partly cover the odontodes situated behind them; their posterior apices are usually broken (Fig. 3S). On the sides of odontodes the surface is finely striated. The basal plate is convex, slightly larger than the crown (Fig. 3S-U). The pores occur in the facets of the basal plate and on the plate around the odontodes, making the surface of the plate uneven.

Histology. Vertical longitudinal (Fig. 4A-G, J), horizontal (Figs 4H, I; 5A, B), and vertical cross-sections (Fig. 5C) were made from both juvenile (Figs 413; 5A, B) and mature (Figs 4A, C-J; 5C, D) scales. Very fine branching dentine tubules occur in the odontodes (Fig. 413, D, G, J). They branch from wider dentine canals (Figs 4B-G, J; 5B), and pulp canals, which in turn are connected with the vascular canal network beneath the odontodes (Figs 4C, E, F, I, J; 513, C). At least one dentine canal extends into each odontode. Vascular canals open on the surface of the basal plate around the odontodes. The basal plate contains tightly packed flattened osteocyte lacunae (Fig. 3C-F, H, J), which give the basal plate structure lamellar appearance. Very fine Sharpey fibre tubules occur in the basal plate (Fig. 5A). One sectioned scale with flat and smooth odontodes (Fig. 5D) has microstructure identical to that of other scales.

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DISCUSSION

Karksilepis gen. nov. is similar to osteostracans by having bone cell cavities in its exoskeleton. Osteostracans have distinct radial structure of peripheral sections of the base of tesserae, which are absent on the basal plate of Karksilepis gen. nov. In Karksilepis gen. nov. the upper layer is comprised of orthodentine, in osteostracans (if present) the equivalent layer consists of mesodentine. Karksilepis gen. nov. differs from osteostracans in having a specific vascular canal network in the basal plate below the odontodes. The ultrasculpture of striation is present in thelodonts, heterostracans, osteostracans, and in Karksilepis parva gen. et sp. nov. The convex basal plate of the scales of Karksilepis gen. nov. is superficially similar to the scale base of acanthodians but their crown sculpture and growth of the scale are different.

The greatest similarities can be found between Karksilepis gen. nov. and some chondrichthyan taxa. Chondrichthyans constitute a very diverse group, with the representatives having very different features. In the elasmobranch chondrichthyan taxa Elegestolepis Karatajute-Talimaa, Kannathalepis Marss & Gagnier, and Ellesmereia Vieth monodontodia are simple, the crown has one pulp cavity, one neck, and one basal canal, while Karksilepis gen. nov. Marss has complex polyodontodia. A possible elasmobranch Seretolepis Karatajute-Talimaa has peculiar lamelliform odontodes on the crown, the base built from aspidine, and basal and neck openings. Knerialepis Hanke & Karatajute-Talimaa has the scales whose odontodes consist of mesodentine and the basal plate of aspidine. Vascular canals are developed between the basal plate and the crown, making Knerialepis similar to Karksilepis, but the sculpture in the former consists of posteriorly fusing wedge-shaped flat odontodes. In Altholepis Karatajute-Talimaa the crown is formed of a great number of narrow anchor-shaped odontodes pointed posteriorly; the scale crown is built from orthodentine and the base--from aspidine. Ivanelepis Karatajute-Talimaa has scales with a few oblong odontodes and neck and basal openings. Arauzia Mader is defined as a possible hybodontiform taxon with rather high scales with upwards directed odontodes of cyclomorial growth and vascular canal openings in the neck and base (Mader 1986). The cladoselachid taxa Iberolepis Mader and Lunalepis Mader have the scale crown and microstructure somewhat resembling those of acanthodians (ibid.). The possible chondrichthyans Frigorilepis Marss et al. and Wellingtonella Marss et al. have monodontode scales without neck canals. In chondrichthyan scales, the neck canals can be present or absent in the same fish, for example, the fin scales of Polymerolepis Karatajute-Talimaa lack neck canals (G. Hanke, pers. comm. 2008) while the scales of other parts of the body have them (see also Karatajute-Talimaa 1998).

Karatajute-Talimaa (1997b) erected the elasmobranch order Lugalepidida based on the characteristics of pectoral and pelvic fins, complex scales in which the odontodes (polyodontodia) of the crown are formed of orthodentine, and the non-growing base of acellular bone tissue; there is no horizontal canal system on the boundary between the crown and the basal plate. According to Karatajute-Talimaa (ibid.), this order differs from Mongolepidida, whose representatives have scales with a growing base and odontodes of lamelline forming longitudinal odontocomplexes of the crown. Karksilepis parva gen. et sp. nov. is externally rather similar to Lugalepis multispinata, which comes from the Emsian(?) and Eifelian of Lithuania, Latvia, Belarus, and Kaliningrad District, Russia (Karatajute-Talimaa 1997b). Both taxa have scales whose crowns consist of numerous spiniform odontodes. Karatajute-Talimaa (1997b) defined such scales in L. multispinata as polyodontodia of synchronomorial type. Nevertheless, these taxa differ in that the upper surface of odontodes of L. multispinata has a longitudinal groove in the medial part, while K. parva gen. et sp. nov. possesses spines with round cross section. Only K. parva gen. et sp. nov. has ultrasculpture of fine striation. The basal plate of the scales of L. multispinata consists of a spongy acellular bone tissue and thin basal lamina, but our new taxon has flattened bone cell lacunae forming the lamellar structure of basal plates.

The scales of Cladodontida also are formed of orthodentine and the basal plate of scales is either of cellular bone tissue (in Cladolepis) or acellular (in Ohiolepis and Protacrodus) (Gross 1973). Karatajute-Talimaa (1997b) noted that some Mongolepidida (Mongolepis, Sodolepis) have an acellular bone developed in the base of scales and some (Teslepis) have a cellular bone with fusiform bone cells; the horizontal system of canals is present in both Mongolepidida and Cladodontida. It would be most appropriate to accommodate Karksilepis gen. nov., having orthodentine in the complex odontodes, horizontal canal network beneath the odontodes in the basal plate, and a cellular bone in the base, within the Cladodontida. Difficulties rise when considering the growth of the crown. In Cladodontida odontodes of approximately the same size were added areally, forming rather regular ring-like zones (Karatajute-Talimaa 1992), while in Karksilepis gen. nov. they occur in irregular rows. As a conclusion, at present we prefer to leave the family and order for Karksilepis parva gen. et sp. nov. Marss open.

RESULTS

Based on the features of scale sculpture, the vascular canal network, and the placement of the canal openings beneath the odontodes in the basal plate, a new chondrichthyan of uncertain order and family, Karksilepis parva gen. et sp. nov. Marss, is established with specimens from the Harma Beds, Burtnieki Regional Stage, Karksi outcrop, Estonia. This is the first time that chondrichthyan dermal scales were discovered in Estonia. Previously only chondrichthyan buccopharyngeal denticles have been documented from the region (Mark-Kurik & Karatajute-Talimaa 2004). The teeth of an elasmobranch from the same outcrop will be discussed in another study.

ACKNOWLEDGEMENTS

We thank J. Valiukevicius for identifying acanthodian scales from Karksi and V. Talimaa (both from Vilnius) for the discussions on the early chondrichthyan scales, and A.Ivanov, St Petersburg, for reading the early version of the manuscript and useful suggestions. The referees Drs Ivan J. Sansom, Birmingham, UK, and Gavin F. Hanke, Victoria, Canada, are thanked for their advice and comments on the original manuscript. SEM images were taken in the Centre of Materials Technology at Tallinn University of Technology (TUT) by Dr. V. Mikli. Thin sections were digitally imaged and all figures improved for the publication by G. Baranov, Institute of Geology at TUT. We thank both for their help. The study was supported financially by the Estonian Science Foundation (grants Nos 5726 and 7334) and by Estonian Target Funding projects SF0140020s08 (for TM) and SF032008OsO7 (for AK).

Received 13 August 2008, accepted 30 September 2008

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Tiiu Marss, Anne Kleesment, and Moonika Niit

Institute of Geology at Tallinn University of Technology, Ehitajate tee 5, 19086 Tallinn, Estonia; Tuu.Marss@gi.ee, kleesmen@gi.ee
Table 1. Distribution of vertebrate microremains in samples from the
Karksi outcrop

                 Vertebrate taxa         Karksi   Karksi   Karksi
                                         02-1     02-2     02-3

Heterostraci     Tartuosteus maximus     xxx      x        x
(Psammosteida)   Pycnosteus              xxx      xx       xx
                 tuberculatus
                 Ganosteus stellatus     x?       x?       --
                 Psammosteus bergi       x?       --       --
                 Psammosteidae gen. et   xx       xxx      xxx
                 sp. indet.
Placodermi                               xx       xx       x

Acanthodii       Diplacanthus            xx       x        --
                 carinatus
                 Diplacanthus sp.        xx       x        x
                 Ptychodictyon           xx       x        xx
                 rimosum
                 Ptychodictyon           xxx      xxx      xxx
                 sulcatum
                 Cheiracanthus           x        x        x
                 longicostatus
                 Cheiracanthus           xx       x        x
                 brevicostatus
                 Cheiracanthus sp.       x        --       x
                 Cheiracanthus           x        --       --
                 sp. nov. A
                 Nostolepis gaujensis    x        xx       --
                 Rhadinacanthus          --       x        --
                 multisulcatus
                 Rhadinacanthus sp.      --       --       x
                 Acanthoides sp. xx      --       --       --
                 Acanthoides sp. A       --       --       --
                 Acanthoides sp. B       --       --       --
                 Acanthoides sp. D       --       --       --
                 Acanthodii gen.         x        xx       xx
                 et sp. indet.
Chondrichthyes   Karksilepis parva       x        x        x
                 gen. et sp. nov.
                 Elasmobranchii          x        --       --
                 gen. et sp. (teeth)
Sarcopterygii                            xxx      xx       xx
Actinopterygii   Cheirolepis sp.         --       x        --

                 Vertebrate taxa         Karksi   Karksi   Karksi
                                         97       90       02-4

Heterostraci     Tartuosteus maximus     x?       x        x
(Psammosteida)   Pycnosteus              xx       x        xx
                 tuberculatus
                 Ganosteus stellatus     x        --       x
                 Psammosteus bergi       x?       --       x?
                 Psammosteidae gen. et   xxx      xx       xxx
                 sp. indet.
Placodermi                               xx       --       xxx

Acanthodii       Diplacanthus            x        x        x
                 carinatus
                 Diplacanthus sp.        --       --       x
                 Ptychodictyon           x        x        x
                 rimosum
                 Ptychodictyon           xx       xx       xx
                 sulcatum
                 Cheiracanthus           x        x        --
                 longicostatus
                 Cheiracanthus           x        x        x
                 brevicostatus
                 Cheiracanthus sp.       x        --       x
                 Cheiracanthus           x        --       --
                 sp. nov. A
                 Nostolepis gaujensis    x        x        x
                 Rhadinacanthus          --       x        --
                 multisulcatus
                 Rhadinacanthus sp.      --       --       x
                 Acanthoides sp. xx      --       --
                 Acanthoides sp. A       --       x        --
                 Acanthoides sp. B       x        x        x
                 Acanthoides sp. D       xx       x        xx
                 Acanthodii gen.         --       --       --
                 et sp. indet.
Chondrichthyes   Karksilepis parva       x        x        x
                 gen. et sp. nov.
                 Elasmobranchii          --       --       x
                 gen. et sp. (teeth)
Sarcopterygii                            xxx     x         xxx
Actinopterygii   Cheirolepis sp.         x       --        --

Findings of vertebrate  microremains: x, rare; xx, common; xxx,
abundant; --not found.
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Author:Marss, Tiiu; Kleesment, Anne; Niit, Moonika
Publication:Estonian Journal of Earth Sciences
Article Type:Report
Geographic Code:4EXES
Date:Dec 1, 2008
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