Geo log i cal Quar terly, 2017, 61 (3): 602–610 DOI: http://dx.doi.org/10.7306/gq.1371
The first microfossil re cord of ichthyofauna from the Naujoji AkmenÅ For ma tion (Lopingian), KarpÅnai Quarry, north ern Lith u a nia
Darja DANKINA1, *, Artur CHAHUD2, Sigitas RADZEVIČIUS1 and Andrej SPIRIDONOV1
1 Vilnius Uni ver sity, De part ment of Ge ol ogy and Min er al ogy, M.K. Čiurlionio 21/27, 03101, Vilnius, Lith u a nia
2 Instituto de GeociÃncias, Departamento de Geologia Sedimentar e Ambiental. - USP. R. do Lago 562, CEP 05508-080, Sao Paulo, Brazil
Dankina, D., Chahud, A., Radzevičius, S., Spiridonov, A., 2017. The first microfossil re cord of ichthyofauna from the Naujoji AkmenÅ For ma tion (Lopingian), KarpÅnai Quarry, north ern Lith u a nia. Geo log i cal Quar terly, 61 (3): 602–610, doi:
10.7306/gq.1371
The as sem blage of rare fish microremains of chon dri chthyans and actinopterygians from Up per Perm ian de pos its in KarpÅnai Quarry in north ern Lith u a nia is de scribed in de tail for the first time. The de pos its are char ac ter ized as re flect ing a rapid phase of de po si tion of the Zechstein Lime stone, in ter preted as on a shal low shelf above storm-wave base in a prox i mal part of the Pol ish-Lith u a nian Zechstein Ba sin. The actinopterygian microremains are rep re sented by var i ous teeth and a few scales from mostly palaeonisciforms and pycnodontiforms. The chondrichthyans are rep re sented by var i ous eusela - chian-type scales and a tooth of ?Helodus sp. The low abun dance and low spe cies di ver sity of the fos sil as sem blages stud - ied may be due to arid palaeoenvironments that caused lo cally re stricted con di tions at this palaeo geo graphi cal lo ca tion.
Key words: chondrichthyans, actinopterygians, Up per Perm ian, Zechstein Ba sin, north ern Lith u a nia.
INTRODUCTION
Late Perm ian chondrichthyans and actinopterygians are known from Aus tra lia (Camp bell and Phuoc, 1983), South Af - rica (Bender, 1999, 2001, 2006), China (Wang et al., 2009), Iran (Hampe et al., 2012), Greece (Argyriou et al., 2017), Rus - sia (Minikh, 1990; Esin, 1993, 1995), and Ger many (Heintz, 1934; Diedrich, 2009).
Un til now only two dis cov er ies of fish microremains have been re ported from the Up per Perm ian of north ern Lith u a nia.
These com prise find ings of a few prob lem atic scales, in ter - preted as Palaeoniscus sp. (Suveizdis, 1963: pl. 13, figs. 8, 9;
Suveizdis, 1975: pl. 19, fig. 5) in the Naujoji AkmenÅ For ma tion in KarpÅnai Quarry. One fish fos sil frag ment, in ter preted as Platysomus sp., has also been found from the Naujoji AkmenÅ For ma tion in the Žalgiriai bore hole (Suveizdis, 1975: pl. 19, fig.
6). These finds were noted with out de tailed de scrip tion or pub - lished il lus tra tion.
The pres ent study of chondrichthyans and actinopterygians is based on iso lated teeth and scales from the Naujoji AkmenÅ For ma tion of KarpÅnai Quarry. Palaeonisciformes (Platyso - midae, Elonichthyidae, Haplolepidae fam i lies) and the Pycno -
dontiformes (Pycnodontidae fam ily) actinopterygians or der have been iden ti fied. The chondrichthyan microremains rep re - sent both the Holocephali and Elasmobranchii.
This study de tails the Up per Perm ian chondrichthyan and actinopterygian tax o nomic and eco log i cal re cords from the north east ern Zechstein Ba sin, which the ter ri tory of the north ern Lith u a nia be longs to. The new data en able a better un der stand - ing of the eco sys tems of the very dy namic Late Perm ian Lopingian Ep och (Kozur, 1995; Clap ham and Bottjer, 2007).
GEOLOGICAL SETTING
Lith u a nia is lo cated in the north east ern part of the Zechstein Ba sin (Kadunas and Raczyński, 2010; Raczyński and Biernacka, 2014) and was lo cated some dis tance away from the main con nec tion be tween the Arc tic Bo real Ocean and the low lat i tude Zechstein Ba sin dur ing the Late Perm ian (SÝrensen et al., 2007; Peryt et al., 2010).
KarpÅnai Quarry is lo cated in north ern Lith u a nia near the Lith u a nian/Lat vian bor der (Fig. 1A). The Naujoji AkmenÅ For ma - tion lime stone (Ca1) is ex posed there, and is a very im por tant min eral re source for the ce ment in dus try in Lith u a nia (Gasiñ - nienÅ and Kadñnas, 1997), in which re spect KarpÅnai Quarry has been stud ied (Kli¹ius, 2006; Kli¹ius and iñraitÅ, 2010).
The Naujoji AkmenÅ For ma tion was first dis tin guished by Suveizdis (1962). Three mem bers (Figs. 1B and 2) were dis tin - guished in the Naujoji AkmenÅ For ma tion (Paškevičius, 1997).
The lower part is char ac ter ized by micritic lime stone (mudstone,
* Corresponding author, e-mail: darja.dankina@gmail.com Received: January 25, 2017; accepted: April 28, 2017; first published online: July 28, 2017
rarely packstone). The mid dle part con sists pre dom i nantly of prox i mal tempestites (grainstone, packstone) with hard lime - stone con cre tions. The up per part is a lime stone show ing mudstone to grainstone tex ture with coated grains. Dolomitic lime stone is found in the high est part of the unit.
Among the en vi ron men tal changes which may have caused sea level fluc tu a tions in the north east ern Zechstein ba sin (Kadñnas, 1997), hu mid ity-arid ity vari a tions have been pro - posed (cf. Biernacka et al., 2005). Cur rently, the quarry ex - poses an al most com plete sed i men tary se quence, with out the low est part of the lower mem ber, and the up per most part of the up per mem ber, of the Naujoji AkmenÅ For ma tion.
The spe cies-rich as sem blage of in ver te brates con sist ing of foraminifers, ostracods, molluscs, brachi o pods, bryo zoans, and echinoderms is dis trib uted through out the Naujoji AkmenÅ For - ma tion (Suveizdis, 1963, 1975), and fau nal di ver sity de creases from the bot tom to the top of the for ma tion (Suveizdis, 1963).
MATERIAL AND METHODS
The new ma te rial co mes from two dif fer ent wall-sec tions (N 56°19.542, E 22°57.219 and N 56°19.503, E 22°57.441 us ing the GPS/WGS84 co or di nate sys tem) in KarpÅnai Quarry. The dis tance be tween these quarry walls is al most 400 m. The height of the ex plor atory quarry walls is ap prox i mately 14 m (Fig. 1B). The sam ples were taken ver ti cally ev ery 2 m from the first site and ev ery 1 m from the sec ond site. As a re sult, 18 sam ples were col lected from both sites. The to tal weight of the sam ples col lected was 210 kg and sam ples var ied from 3 to 22 kg (Ta bles 1 and 2).
All sam ples were dis solved with buf fered 10% for mic acid (CH2O2) for the dolomitized de pos its, and with buf fered 10%
ace tic acid (CH3COOH) for the lime stones (Jeppsson et al., 1999). The res i due was nat u rally dried and sieved from 0.2 to 0.063 mm to ex tract the ex tremely tiny fish microremains and Fig. 1. Lo ca tion of KarpÅnai Quarry
A – ex tent of the Zechstein Lime stone in the east ern part of the South ern Perm ian Ba sin in Cen tral Eu rope (af ter Peryt et al., 2012);
B – KarpÅnai Quarry stra tig ra phy: lower part (L) – micritic lime stone, mid dle part (M) – prox i mal tempestites, up per part (U) – dolomitic lime stone
ease the pick ing pro cess. The microremains were ex tracted onto microslides. A se lected num ber of the microremains was in ves ti gated and pho to graphed with a scan ning elec tron mi cro - scope (SEM) at the Na ture Re search Cen tre (Vilnius, Lith u a - nia) and at the Uni ver sity of Va len cia (Va len cia, Spain).
All the ma te rial is stored in the Geo log i cal Mu seum of the In - sti tute of Geosciences Vilnius Uni ver sity (Lith u a nia).
ACTINOPTERYGIAN FAUNA
The actinopterygians are rep re sented by their iso lated teeth and scales (Ta bles 1 and 2) and are iden ti fied as Palaeo - nisciformes and Pycnodontiformes.
94 teeth and 8 scales are as signed to the Palaeo nis - ciformes or der (Fig. 3F–K) on the ba sis of a com par i son with Palaeonisciformes-re lated ma te rial (Štamberg, 2016: fig. 4–6) and its de scrip tion (Esin, 1990). Most of the teeth are low, not ex ceed ing 1.2 mm in length, and the widths of their bases reached 0.3 mm (Fig. 3G–I). The teeth are con i cal, straight, and smooth, with acrodin caps (Fig. 3H, J, K), or miss ing caps (Fig.
3G, I). The dis tinct acrodin cap is cir cu lar in api cal view, reach - ing a max i mum di am e ter of 0.18 mm, while in la bial or lin gual view, the acrodin cap is tri an gu lar in out line and has a length of 0.05 mm. The cor pus of these rostral teeth is sculp tured, with the ex cep tion of the acrodin apex, with shal low ridges uni formly di rected from the acrodin apex to the base of the tooth (Fig. 3H).
Some teeth have no dis tinct sculp ture, though micro - tubercles are well-de vel oped and cover the en tire pre served tooth sur face. These microtubercles are nar row and ver ti cally elon gated, with oblique rows (Fig. 3G, I–K). Most scales have a rhomboidal, and in places more elon gated, drop-like (Fig. 3F) shape with a poorly pre served outer sur face. The sur face is smooth, with no dis tinct or na men ta tion. The base is flat and thick, with no keel. These scales reach a max i mum length of 1.1 mm and have a width of 0.6 mm. The teeth and scales stud ied from KarpÅnai Quarry are poorly pre served. Most teeth have a dis - tinct or na ment, com pa ra ble with microsculpture of sim i lar teeth (tamberg, 2016). It is dif fi cult to iden tify Palaeoniscus sp. based solely on the mor phol ogy of iso lated scales, as there is a lack of com par a tive in - for ma tion about the scales (Esin, 1990).
One tooth (VU-ICH-KAR-004) has been as - signed to “Elonichthys” sp. (Fig. 3D). The tooth is slen der and sharp-pointed in shape, with a length of 0.6 mm, and the width of its base is 0.15 mm. The dis tinct acrodin apex has a length of 0.1 mm (Fig.
3D2). The microtubercles are well-de vel oped. They cover the whole sur face of the tooth, with the ex cep - tion of the acrodin apex. The prox imo-dis tally micro - tubercles are nar rower, more elon gated, and blend to gether (Fig. 3D3). The tooth has been as signed to Elonichthys based on com par i son to the iso lated tooth of “Elonichthys” sp. from the Black Shale Ho ri - zon of the Syřenov For ma tion, Up per Car bon if er ous in the Krkonoše Piedmont Ba sin (Štamberg, 2016:
fig. 10A–C) which bears sim i lar tu ber cles.
Two iso lated teeth from the Platysomidae fam ily (Fig. 3A, B) were found in the re gion un der study. One tooth (VU-ICH-KAR-001) is con i cal and de void of or - na men ta tion (Fig. 3A). It reaches 1.2 mm in height, whereas the di am e ter at the base does not ex ceed 0.7 mm. One crush ing-type tooth (VU-ICH-KAR-002) is oval in the occlusal-lat eral view, which reaches a di am e ter of 0.5 mm in api cal view and a height of 0.6 mm (Fig. 3B1). The occlusal sur face is con vex (Fig. 3B2). The as so ci a tion of con i cal and broad, crush ing-type teeth is typ i cal of Platysomus. There is a gra da tion be tween rounded and pointed teeth (John son and Zidek, 1981). Their mor phol ogy con forms with the de scrip tions of platysomid fish tooth plates from the up per Clyde For ma tion, Up - per Pa leo zoic (John son and Zidek, 1981: text-fig. 3D, E).
Iso lated scales (VU-ICH-KAR-005) of the Haplolepidae fam - ily (Fig. 3E) are small, reach ing up to 0.8 mm in length and 0.6 mm in width. The scales are rhom boid with four sharp edges (Fig. 3E1, E2). A layer of ganoine tis sue cov ers the poorly pre - served outer sur face. The sur face has weakly pro nounced sin u - ous par al lel grooves (Fig. 3E1). This type of scale is prob a bly lo - cated at the pos te rior part of the trunk where the fish body ta pers in the cau dal fin area (Trinajstic, 1977). The bases of the scales are thick and con vex, with no keel. The iso lated rhombic scales are as signed to the Haplolepidae fam ily on the ba sis of a com - par i son with more com plete ma te rial (Westoll, 1944). A sim i lar rhomboidal scale from a haplolepid with a ganoine-cov ered outer sur face and sin u ous nar row grooves was found in the Rader Lime stone Mem ber of the Bell Can yon For ma tion, Capitanian in West Texas, USA (Ivanov et al., 2015: pl. 2, fig. 15).
Two iso lated teeth from the Pycnodontidae fam ily (Fig. 3C) were found at the site un der study. The teeth (VU-ICH -KAR - -003) are cir cu lar in api cal view, reach ing a max i mum di am e - ter of 1.2 mm (Fig. 3C3). These teeth are low, not ex ceed ing 0.4 mm in height. The sur face of the acrodin caps is densely 604 Darja Dankina, Artur Chahud, Sigitas Radzevičius and Andrej Spiridonov
T a b l e 1 KarpÅnai ex po sure I
No. Height
(m) Spec i men Weight
(kg) Actinopterygia
Chondrichthyan Scale Tooth Scale Tooth
1 12.0–14.0 KAR-7 4,4 0 1 1 0
2 10.0–12.0 KAR-6 5,25 0 2 6 0
3 8.0–10.0 KAR-5 3,70 0 0 4 0
4 6.0–8.0 KAR-4 12,04 0 1 1 0
5 4.0–6.0 KAR-3 7,86 0 9 7 0
6 2.0–4.0 KAR-2 3,10 1 21 11 0
7 0.0–2.0 KAR-1 9,58 1 7 6 0
To tal: 45.93 2 41 36 0
T a b l e 2 KarpÅnai ex po sure II
No. Height
(m) Spec i men Weight (kg)
Actinopterygian Chondrichthyan Scale Tooth Scale Tooth
1 13.0–14.0 KAR-18 14,2 0 1 2 0
2 12.0–13.0 KAR-17 22,4 0 3 1 0
3 11.0–12.0 KAR-16 11,6 0 0 0 0
4 10.0–11.0 KAR-15 12,9 0 5 10 0
5 9.0–10.0 KAR-14 19,3 2 10 22 0
6 8.0–9.0 KAR-13 12,9 0 8 17 0
7 7.0–8.0 KAR-12 13,2 0 0 1 1
8 6.0–7.0 KAR-11 6,8 0 0 3 0
9 5.0–6.0 KAR-10 18 0 16 2 0
10 4.0–5.0 KAR-9 19,3 1 5 14 0
11 3.0–4.0 KAR-8 14 4 8 6 0
To tal: 164.6 7 56 78 1
or na mented, with ir reg u lar, ra di at ing, and anastomosing ridges (Fig. 3C1). These ridges do not join in the mid dle of the cap tops, leav ing a small area in the cen ter (Fig. 3C2). The low pro file and cir cu lar out line of these teeth sug gest pycnodont af fin i ties. A sim i lar cir cu lar tooth with ra di at ing merg ing ridges has been il lus trated and iden ti fied as cf. Anomoeodus from the Sao Khua For ma tion of the Cre ta ceous at Phu Phan Thong in Thai land (Cavin et al., 2009: fig. 9F). Also, the teeth stud ied were found to have mor pho log i cal sim i lar i ties to Pycnodo - ntiformes indet. from the Lower Cre ta ceous in Tu ni sia (Cuny et al., 2010: fig. 4-1a).
CHONDRICHTHYAN FAUNA
The chondrichthyans are rep re sented by iso lated re mains and iden ti fied as euselachian type scales and a ?Helodus tooth (Ta bles 1 and 2).
The tooth stud ied (VU-ICH-KAR-012) (Fig. 4A) mea sures 2.6 mm mesio-dis tally and is 1.5 mm high, the root be ing ap - prox i mately half the height of the crown (Fig. 4A1). The root is per fo rated by nu mer ous, ran domly dis trib uted fo ram ina on the la bial and lin gual sides (Fig. 4A3). It com prises a sin gle, some -
what ir reg u lar row of full and crushed dif fer ent-sized ca nal open ings be low the crown. The tooth crown is de void of or na - men ta tion. It is oval with a smooth edge and one nar rower sharp point (Fig. 4A2). The tooth has been pu ta tively as signed to the Helodontidae fam ily. The gen eral shape of this tooth is com pa - ra ble to the Helodus-like tooth de scribed by East man (1903: pl.
2, fig. 14) and to a de scrip tion of a Helodus tooth in Stahl (1999). How ever, this tooth is also sim i lar to that of a hybodont Lissodus sp., from the Akasaka Lime stone For ma tion, the Mid - dle Perm ian in Cen tral Ja pan (Yamagishi and Fujimoto, 2011:
fig. 2E, F) and to a Polyacrodus teeth from Early Perm ian strata in Texas (John son, 1981). There fore, it is not pos si ble to be sure that the tooth stud ied def i nitely be longs to the Helodus sp.
The 114 iso lated euselachian type scales (Fig. 4B–K) be - long to sev eral morphotypes based on the crown mor phol ogy.
Morphotype A (Fig. 4B, D) is rep re sented by typ i cal placoid scales with a flat, thick, smooth, or some times po lyg o nal crown, and a high base. In most cases, the crown has a hor i zon tal ex - ter nal sur face and ver ti cal lat eral sides. The edges of the crown are rounded and lo cally un du late with tiny, short ridges. The crown and base bound ary area is dis tinct with a well-de vel oped and pre served neck. The base has nu mer ous fo ram ina of vas - cu lar ca nals. In places the base is higher and wider than the crown, and has an al most ver ti cal lat eral and flat basal sur face.
Fig. 2A – strati graphic sub di vi sions of the Late Perm ian suc ces sion in the Lith u a nia and Kaliningrad re gion (Kadunas and Raczyński, 2010; Raczyński and Biernacka, 2014); B – finds of chondrichthyan and actinopterygian microfossils
606 Darja Dankina, Artur Chahud, Sigitas Radzevičius and Andrej Spiridonov
Fig. 3. Actinopterygian scales and teeth from the Naujoji AkmenÅ For ma tion, KarpÅnai Quarry
A – Platysomidae indet. teeth, lat eral view (VU-ICH-KAR-001); B1 – Platysomidae indet. teeth, occlusal-lat eral view, B2 – occlusal sur face (VU-ICH-KAR-002); C1 – ?Pycnodontidae indet. teeth, with ra di at ing ridges, C2 – a cen ter of the cap top, C3 – api cal view (VU-ICH-KAR-003);
D1 – ?"Elonichthys" sp. tooth, lat eral view, D2 – the dis tinct acrodin apex, D3 – the elon gated prox imo-dis tally microtubercles (VU-ICH-KAR-004); E1 – ?Haplolepidae indet. scale, poorly pre served outer sur face, E2 – in ner sur face (VU-ICH-KAR-005); G1 – Palaeonisciformes indet. teeth, lat eral view, G2 – nar row, ver ti cally elon gated microturbercles (VU-ICH-KAR-007)
Fig. 4. Chondrichthyan scales and tooth from the Naujoji AkmenÅ For ma tion
A – ?Helodontidae indet. tooth: A1 – lat eral view, A2 – occusal view, A3 – lat eral/basal view (VU-ICH-KAR-012); B, D – euselachian type scales of morphotype A: B1 – crown view, B2 – an te rior view, B3 – basal view (VU-ICH-KAR-013), D – crown view (VU-ICH-KAR-014); C, E–K – euselachian type scales of morphotype B: C – an te rior view (VU-ICH-KAR-015), E – crown view (VU-ICH-KAR-016), F – crown view (VU-ICH-KAR-017), G – lat eral view (VU-ICH-KAR-018), H – postero-basal view (VU-ICH-KAR-019), I – postero-basal view, (VU-ICH- KAR-020), J – crown view (VU-ICH-KAR-021), K – crown view (VU-ICH-KAR-022)
608 Darja Dankina, Artur Chahud, Sigitas Radzevičius and Andrej Spiridonov
The round ish scales range in di am e ter to 0.6 mm (Fig. 4B), while the po lyg o nal crown is 1.0 mm in width and 0.8 mm in length (Fig. 4D). The morphotype B scales (Fig. 4C, E–K) have a drop-like, elon gated, or rarely oval crown (Fig. 4J), a well-de - vel oped neck, which is smaller, or some times big ger, than the crown base. The crown is strongly in clined and the pointed pos - te rior edge over tops be neath the blunt an te rior part. The slightly con vex ex ter nal sur face of the crown is smooth (Fig. 4E, G), or in places has dis tinct me dial ridges. The me dial ridge var ies from be ing short, reach ing an te rior edge (Fig. 4C, K) to be ing wide and long, reach ing the pos te rior edge (Fig. 4F). The scale neck is nar row, or in some in stances wide (Fig. 4H), low in the an te rior part and high pos te ri orly. The base has a con cave (Fig.
4I) or flat basal sur face. The crowns of scales are small, from 0.2–0.8 mm in length, and 0.4 mm in width. A po lyg o nal crown sim i lar to that from morphotype A (Fig. 4D) has been de scribed by Reif (1978: fig. 9C) from the Lower Muschelkalk, the Mid dle Tri as sic in Cen tral Ger many, and as signed to the Hybodontidae fam ily. Al though, the an te rior part of the Hybodus scale, as pre - sented by Reif (1978), has a dis tinct con stric tion. How ever, the hybodontids have scales very sim i lar to the placoid scales of the euselachians (Reif, 1978). More over, the sim i lar crown with un du lat ing ridges (Fig. 4B) was named a Jannasa korni placoid scale (Brandt, 1996) from the Zechstein Lime stone of Ger - many. Scales sim i lar to morphotype B scales have been il lus - trated and de scribed from the Mid dle Perm ian in West Texas, USA as euselachian scales by Ivanov et al. (2013: fig. 5A, B).
Manzanares et al. (2014: fig. 1a) named sim i lar drop-like scales from the Mid dle Tri as sic in Spain as Glabrisubcorona-type hybodontiform scale based on their his tol ogy. Many scales sim - i lar to morphotype A and morphotype B based on their crown mor phol ogy were found in Mid dle and Late Tri as sic strata in Brit ish Co lum bia, Can ada and de scribed as elasmobranch scales (Johns, 1996).
DISCUSSION
The ver te brate as sem blage from the Naujoji AkmenÅ For - ma tion in KarpÅnai Quarry com prises actinopterygian and chon drichthyan ma te rial of low abun dance and Late Perm ian age. The actinopterygian re mains are rep re sented by teeth and scales from mostly palaeonisciforms and pycnodontiforms, while the chondrichthyans are rep re sented by var i ous eusela - chian -type scales and a ?Helodus sp. tooth.
The Late Perm ian palaeoniscoids were widely dis trib uted geo graph i cally and oc curred in many en vi ron ments. For ex am - ple, rep re sen ta tives of this taxon oc cur in lo cal i ties rang ing from Aus tra lia and South Af rica (Camp bell and Phuoc, 1983; Ben - der, 1999, 2001, 2006); to Eu ro pean Rus sia (Minikh, 1990;
Esin, 1995). Whole Platysomidae skel e tons and some iso lated frag ments have been found in the Up per Kazanian Stage, the Mid dle Perm ian, and Up per Perm ian of the East Eu ro pean Plat form of Rus sia (Solodukho, 1951; Esin, 1993); well-pre - served Platysomus sp. and Palaeoniscum sp. skel e tons have been found in the Lopingian Stage, the Up per Perm ian in north - west Ger many (Diedrich, 2009). Elonichthys sp. skel e tons and skull frag ments came from Permo-Car bon if er ous lakes strata in the Czech Re pub lic and in the Saar-Nahe Ba sin in south-west Ger many (Schindler, 1993; Zajíc, 2004, 2007; Štamberg, 2014); some Elonichthys sp. microremains have been found in Perm ian de pos its in Nor way (Heintz, 1934). Haplolepis sp. re - mains are known from the Mid dle Perm ian of West Texas, USA (Ivanov et al., 2015).
Four main or ders of chondrichthyans: Hybodontiformes, Xena canthiformes, Symmoriiformes, and Ctenacanthiformes are rep re sented in Mid dle and Late Perm ian bas ins in East ern Eu rope (Minikh and Minikh, 2009). More over, euselachian scales are known from the Kazanian Stage, Mid dle Perm ian, of Rus sia (Ivanov and Lebedev, 2014). Also, Helodus sp. frag - ments have been found in Perm ian de pos its of West Aus tra lia (Teichert, 1943).
A more de tailed tax o nomic anal y sis of the fish fauna in the KarpÅnai Quarry needs ad di tional and well-pre served ma te rial, al low ing microstructural and histological in ves ti ga tion. That might then help Up per Perm ian strati graphic cor re la tion in the Pol ish-Lith u a nian part of the Zechstein Ba sin.
The rar ity of these fish microremains can be ex plained in sev - eral ways. The Zechstein Ba sin had some sim i lar i ties with the palaeoenvironmental sit u a tion in the Mio cene Med i ter ra nean Sea dur ing the Late Mio cene Sa lin ity Cri sis. The Med i ter ra nean Sea was at least par tially iso lated from the At lan tic Ocean, and flow of less sa line wa ter from the ocean was re stricted. As a re - sult, the east ern part of the ba sin was hypersaline and highly re - stricted (Blanc, 2006). The same ef fects may have af fected to some ex tent the Lith u a nian part of the ba sin, from its dis tal lo ca - tion from the Arc tic Bo real Ocean dur ing the Late Perm ian (SÝrensen et al., 2007). In creased sa lin ity may be one po ten tial lim it ing fac tor for the dis tri bu tion of fish. Also, the dis ap pear ance of the thermocline in coastal ar eas of upwelling may cause ex - treme ma rine fer til ity and an oxia pro mot ing the de vel op ment of bi tu mi nous sed i ments (Brongersma-Sanders, 1971). The lime - stone and dolostone sam ples stud ied were en riched with bi tu - men, es pe cially, in the mid dle mem ber in KarpÅnai Quarry. High phytoplankton pro duc tiv ity may have caused eutro phication, to re sult in the de ple tion of dis solved ox y gen in the wa ter.
More over, Lith u a nia’s coastal zone was much wider than those of the south ern parts of the ba sin (e.g., in SW Po land) as a re sult of pro longed ero sion of the plat form, and low top o - graphic re lief across a large ter ri tory (Raczyński and Biernacka, 2014). This pos si bly re duced cir cu la tion, which could have re - sulted in re stricted high-tem per a ture and low-ox y gen con di - tions, which also did not fa vour con di tions for ver te brates. All these fac tors may have con trib uted to the low di ver sity and abun dance of the fos sil fish in the area stud ied.
CONCLUSIONS
Study of the KarpÅnai Quarry sec tions yielded chondri - chthyan and actinopterygian as sem blages, usu ally ex tremely rare in the Up per Perm ian of north ern Lith u a nia. Large sam ples of tens to hun dreds of kilogram of rock are needed to yield suf fi - cient sam ple sizes to re veal the di ver sity of the microvertebrate re mains.
The poor tax o nomic di ver sity of fish fauna in the KarpÅnai lo cal ity may be ex plained by the for mer pres ence of re stricted, high-sa lin ity en vi ron ments, due to the gen er ally flat coastal zone of the northeast ern shore of the Zechstein Sea, and the very dis tal lo ca tion from the Bo real Ocean.
Ac knowl edge ments. We would like to ex press our cor dial thanks to A. Ivanov (St. Pe ters burg State Uni ver sity, Rus sia), and V. KarajñtÅ-Talimaa (Vilnius Uni ver sity, Lith u a nia), who helped with the iden ti fi ca tion of our ma te rial. Many thanks to A. Ivanov for his use ful pa pers on the Perm ian ichthyofauna and to P. Raczyński (Uni ver sity of Wroclaw, Po land), for his help ful re views on the stra tig ra phy of the area stud ied, and to
L. Šiliau skas (Na ture Re search Cen ter, Lith u a nia) and E. Manzanares (Uni ver sity of Va len cia, Spain) for tak ing SEM mi cro graphs. We would also like to thank A. Ivanov and M.
Ginter (Uni ver sity of War saw, Po land) for their many cru cial re - marks which sig nif i cantly im proved the ar ti cle. This re search
was car ried out by the Open Ac cess to the re search in fra struc - ture of the Na ture Re search Cen tre (Vilnius) un der the Lith u a - nian open ac cess net work ini tia tive.
REFERENCES
Argyriou, T., Romano, C., Carrillo-BriceÔo, J.D., Brosse, M., Hofmann, R., 2017. The old est re cord of gnathostome fos sils from Greece: Chondrichthyes from the Lopingian of Hy dra Is - land. Palaeontologia Electronica, 20: 1–9.
Bender, P.A., 1999. First doc u men ta tion of sim i lar Late Perm ian actinopterygian fish from Aus tra lia and South Af rica. Re cords of the West ern Aus tra lian Mu seum, 57: 183–189.
Bender, P., 2001. A new actinopterygian fish spe cies from the Late Perm ian Beau fort Group, South Af rica. Palaeontologia Africana, 37: 25–40.
Bender, P., 2006. A new deep-bod ied Late Perm ian actinopterygian fish from the Beau fort Group, South Af rica. Palaeontologia Africana, 41: 7–22.
Biernacka, J., Borysiuk, K., Raczyński, P., 2005. Zechstein (Ca1) lime stone-marl al ter na tions from the North-Sudetic Ba sin, Po - land: depositional or diagenetic rhythms? Geo log i cal Quar terly, 49 (1): 1–14.
Blanc, P.L., 2006. Im proved mod el ling of the Messinian Sa lin ity Cri - sis and con cep tual im pli ca tions. Palaeo ge ogra phy, Palaeo - climatology, Palaeo ec ol ogy, 238: 349–372.
Brandt, S., 1996. Janassa korni (Weigelt) – Neubeschreibung eines petalodonten Elasmobranchiers aus dem Kupferschiefer und Zechsteinkalk (Perm) von Eisleben (Sachsen-Anhalt).
Paläonto logische Zeitschrift, 70: 505–520.
Brongersma-Sanders, M., 1971. Or i gin of ma jor cyclicity of eva - porites and bi tu mi nous rocks: an actualistic model. Ma rine Ge - ol ogy, 11: 123–144.
Camp bell, K.S.W., Phuoc, L.D., 1983. A Late Perm ian actino - pterygian fish from Aus tra lia. Palae on tol ogy, 26: 33–70.
Cavin, L., Deesri, U., Suteethorn, V., 2009. The Ju ras sic and Cre - ta ceous bony fish re cord (Actinopterygii, Dipnoi) from Thai land.
Geo log i cal So ci ety Spe cial Pub li ca tions, 315: 125–139.
Clap ham, M.E., Bottjer, D.J., 2007. Perm ian ma rine paleoecology and its im pli ca tions for large-scale de coup ling of brachi o pod and bi valve abun dance and di ver sity dur ing the Lopingian (Late Perm ian). Palaeo ge ogra phy, Palaeoclimatology, Palaeo ec ol - ogy, 249: 283–301.
Cuny, G., Cobbett, A.M., Meunier, F.J., Benton, M.J., 2010. Ver te - brate microremains from the Early Cre ta ceous of south ern Tu ni - sia. Geobios, 43: 615–628.
Diedrich, C.G., 2009. A coelacanthid-rich site at Hasbergen (NW Ger many): taphonomy and palaeoenvironment of a first sys tem - atic ex ca va tion in the Kupferschiefer (Up per Perm ian, Lopingian). Palaeobiodiversity and Palaeoenvironments, 89:
67–94.
East man, C.R., 1903. Car bon if er ous fishes from the Cen tral West - ern States. Bul le tin of the Mu seum of Com par a tive Zo ol ogy, 39:
161–226.
Esin, D.N., 1990. The scale cover of Amblypterina costata (Eich - wald) and the palaeoniscid tax on omy based on iso lated scales (in Rus sian). Paleontological Jour nal, 2: 90–98.
Esin, D.N., 1993. New spe cies of deep-bod ied actinopterygians (Platysomidae) from the Up per Perm ian of the East Eu ro pean Plat form (in Rus sian). Paleontological Jour nal, 3: 166–171.
Esin, D.N., 1995. Ontogenetic de vel op ment of the squamation in some palaeoniscoid fishes. Bul le tin du Muséum Na tional d’Histoire Naturelle, 17: 227–234.
GasiñnienÅ, V.E., Kadñnas, V., 1997. Lietuvos viršutinio permo klinties pramoninÅ charakteristika (in Lith u a nian). Geologija, 22:
50–55.
Gailius, R., Grigelis, A., Jankauskas, T., Juodkazis, V., Juozapavičius, G., Kadñnas, V., Katinas, V., Klimas, A., Kondratas, A., KondratienÅ, O., Linčius, A., Mikalauskas, V., Narbutas, V., Paškevičius, J., Sakalauskas, K., Suveizdis, P., Tarvydas, R., Vonsavičius, V., 1994. Lietuvos geologija (in Lith u a nian). Mokslo ir Enciklopedij÷ Leidykla, Vilnius.
Hampe, O., Hairapetian, V., Dorka, M., Witzmann, F., Akbari, A.M., Korn, D., 2012. A first late Perm ian fish fauna from Baghuk Moun tain (Neo-Tethyan shelf, cen tral Iran). Bul le tin of Geosciences, 88: 1–20.
Heintz, A., 1934. Fischreste aus dem Unterperm Norwegens. Norsk Geologisk Tidsskrift, 14: 176–194.
Ivanov, A.O., Nestell, G.P., Nestell, M.K., 2013. Fish as sem blage from the Capitanian (Mid dle Perm ian) of the Apache Moun tains, West Texas, USA. New Mex ico Mu seum of Nat u ral His tory and Sci ence, Bul le tin, 60: 152–160.
Ivanov, A.O., Lebedev, O.A., 2014. Perm ian chondrichthyans of the Kanin Pen in sula, Rus sia. Paleontological Jour nal, 48:
1030–1043.
Ivanov, A.O., Nestell, M.K., Nestell, G.P., 2015. Mid dle Perm ian fish microremains from the Early Capitanian of the Guadalupe Moun tains, West Texas, USA. Micropaleontology, 61: 301–312.
Jeppsson, L., Anehus, R., Fredholm, D., 1999. The op ti mal ac e - tate buf fered ace tic acid tech nique for ex tract ing phos phatic fos - sils. Journa of Pa le on tol ogy, 73: 964–972.
Johns, M.J., 1996. Di ag nos tic pedicle fea tures of Mid dle and Late Tri as sic elasmobranch scales from north east ern Brit ish Co lum - bia, Can ada. Micropaleontology, 42: 335–350.
John son, G.D., 1981. Hybodontoidei (Chondrichthyes) from the Wich ita-Al bany Group (Early Perm ian) of Texas. Jour nal of Ver - te brate Pa le on tol ogy, 1: 1–41.
John son, G.D., Zidek, J., 1981. Late Pa leo zoic phyllodont tooth plates. Jour nal of Pa le on tol ogy, 55: 524–536.
Kadñnas, V., 1997. Sedimentacini÷ proces÷ itaka viršutinio permo klinties klodo storiui ir mineralinei sudÅčiai ŠiaurÅs Lietuvoje (in Lith u a nian). Geologija, 22: 36–42.
Kadñnas, V., 2001. Lietuvos permo halogeninÅ formacija: litologija, geochemija, naudingosios iškasenos (in Lith u a nian). Geolo - gijos institutas, Vilnius.
Kadunas, V., Raczyński, P., 2010. Lith u a nia and Lat via – the east - ern most part of the South ern Perm ian Ba sin. In: Pe tro leum Geo - log i cal At las of the South ern Perm ian Ba sin Area (eds. J.C.
Doornenbal and A.G. Stevenson): 141–142. EAGE Pub li ca tions b.v. (Houten).
Kličius, J., 2006. AkmenÅs rajono KarpÅn÷ klinties telkinio naujo ploto detalios žvalgybos ataskaita (in Lith u a nian). Gelmi÷
Tyrimai, Vilnius.
Kličius, J., ČiñraitÅ, K., 2010. AkmenÅs rajono KarpÅn÷ klinties telkinio naujo ploto detalios žvalgybos ataskaita (in Lith u a nian).
Gelmi÷ Tyrimai, Vilnius.
Kozur, H., 1995. Perm ian cono dont zonation and its im por tance for the Perm ian strati graphic stan dard scale. Geologisch -Paläonto - logische Mitteilungen Innsbruck, 20: 165–205.
Manzanares, E., Plá, C., Martínez-Pérez, C., Rasskin, D., Botella, H., 2014. The enameloid microstructure of euselachian (Chon - drichthyes) scales. Paleontological Jour nal, 48: 1060–1066.
Minikh, A.V., 1990. Novyi paleonisk iz pozdney permi vostochno - -evropeyskoy platformy (in Rus sian). Paleontologicheskiy Zhur - nal, 3: 71–76.
Minikh, A.V., Minikh, M.G., 2009. Osnovnye sobytiya v razvitii sredne i pozdnepermskoy ikhtiofauny Vostochnoy Evropy (in Rus sian). Nauchnaya Kniga, 141–157.
Paškevičius, J., 1997. The Ge ol ogy of the Bal tic Re pub lics. Vilnius Uni ver sity, Geo log i cal Sur vey of Lith u a nia, Vilnius.
Peryt, T.M., Geluk, M.C., Mathiesen, A., Paul, J., Smith, K., 2010.
Zechstein. In: Pe tro leum Geo log i cal At las of the South ern Per - m ian Ba sin Area (eds. J.C. Doornenbal and A.G. Stevenson):
123–147. EAGE Pub li ca tions b.v. (Houten).
Peryt, T.M., Durakiewicz, T., Kotarba, M.J., Oszczepalski, S., Peryt, D., 2012. Car bon iso tope stra tig ra phy of the basal Zech - stein (Lopingian) strata in North ern Po land. Geo log i cal Quar - terly, 56 (2): 285–298.
Raczyński, P., Biernacka, J., 2014. Zechstein in Lith u a nia-Lat via bor der re gion. Geologija, 2: 57–62.
Reif, W.E., 1978. Types of morphogenesis of the der mal skel e ton in fos sil sharks. Paläontologische Zeitschrift, 5: 110–128.
Schindler, T., 1993. “Elonichthys” pal a tines n. sp., a new spe cies of actinopterygians from the Lower Perm ian of the Saar-Nahe Ba - sin (SW-Ger many). New Re search on Permo-Car bon if er ous Fau nas. Pollichia-Buch, 29: 67–81.
Solodukho, M.G., 1951. Nakhodki predstaviteley sem. Platyso - midae v verkhnekazanskikh otlozheniyakh okrestnostey d.
Pechishchi (Tatarskaya ASSR) (in Rus sian). Uchyonnye Zapisi kazanskogo Gosudarstvennogo Universiteta im. V.I. Ul’yanova - -Lenina, 111: 157–159.
SÝrensen, A.M., HDkansson, E., Stemmerik, L., 2007. Fau nal mi - gra tion into the Late Perm ian Zechstein Ba sin – ev i dence from bryo zoan palaeobiogeography. Palaeo ge ogra phy, Palaeo - clima tology, Palaeo ec ol ogy, 251: 198–209.
Stahl, B.J., 1999. Chondrichthyes III. Holocephali. In: Hand book of Paleoichthyology (ed. H.P. Schultze). Friedrich Pfeil, München.
Suveizdis, P., 1962. Stratigrafiya permskikh otlozheniy Polsko - -Litovskoy sineklizy po novym paleontologicheskim materialam (in Rrussian). Moksliniai Pranešimai: Geologija Geografija, 14:
5–32.
Suveizdis, P., 1963. Verkhnepermskie otlozheniya Polsko -Litovskoj sineklizy (in Rus sian). Voprosy Geologii Litvy, Vilnius.
Suveizdis, P. (ed.), 1975. Perm ian de pos its of the Bal tic re gion (stra tig ra phy and fauna) (in Rus sian with Eng lish sum mary).
Lietuvos Geologijos Mokslinio Tyrimo Institutas, Trudy, Vilnius.
Štamberg, S., 2014. Dis cov ery of skel e tal frag ments of a large am - phib ian and other Perm ian fauna from lo cal ity Arnultovice in the Krkonoše Piedmont Ba sin (in Czech). Geoscience Re search Re ports, 47: 94–97.
Štamberg, S., 2016. Actinopterygians of the Stephanian sed i ments of the Krkonoše Piedmont Ba sin (Bo he mian Mas sif) and their palaeobiogeographic re la tion ship. Bul le tin of Geosciences, 91:
169–186.
Teichert, C., 1943. Bradyodont sharks in the Perm ian of West ern Aus tra lia. Amer i can Jour nal of Sci ence, 243: 543–552.
Trinajstic, K., 1977. Scale mor phol ogy of the Late De vo nian palaeo niscoid Moythomasia durgaringa Gar di ner and Bartram 1977. Al che rin ga, 23: 9–19.
Wang, N.Z., Zhang, X., Zhu, M., Zhao, W.J., 2009. A new ar tic u - lated hybodontoid from Late Perm ian of north west ern China.
Acta Zoologica, 90: 159–170.
Westoll, T.S., 1944. The Haplolepidae, a new fam ily of Late Car bon - if er ous bony fishes. Bul le tin of the Amer i can Mu seum of Nat u ral His tory, 83: 1–122.
Yamagishi, H., Fujimoto, T., 2011. Chondrichthyan re mains from the Akasaka Lime stone For ma tion (Mid dle Perm ian) of Gifu Pre - fec ture, Cen tral Ja pan. Bul le tin of the Kanagawa Pre fec tural Mu seum, Nat u ral Sci ence, 40: 1–6.
Zajíc, J., 2004. Ver te brate biozonation of the Permo-Car bon if er ous lakes of the Czech Re pub lic – new data. Acta Musei Reginaehradecensis S.A., 30: 15–16.
Zajíc, J., 2007. Car bon if er ous Fauna of the Krkonoše Piedmont Ba - sin. Acta Musei Reginaehradecensis S.A., 32: 11–16.
610 Darja Dankina, Artur Chahud, Sigitas Radzevičius and Andrej Spiridonov
