GeolCarp_Vol70_No3_209

GEOLOGICA CARPATHICA

, JUNE 2019, 70, 3, 209–221

doi: 10.2478/geoca-2019-0012

www.geologicacarpathica.com

Graptolite turnover and δ

org

excursion in the upper

Wenlock shales (Silurian) of the Holy Cross

Mountains (Poland)

SIGITAS RADZEVIČIUS



, PAWEŁ RACZYŃSKI

, MARIUS UŽOMECKAS

AUDRIUS NORKUS

and ANDREJ SPIRIDONOV

1, 3

Department of Geology and Mineralogy, Vilnius University, M.K. Čiurlionio 21/27, LT-03101, Vilnius, Lithuania;



sigitas.radzevicius@gf.vu.lt

Institute of Geological Sciences, University of Wrocław, Pl. Maksa Borna 9, Wrocław 50-205, Poland

Laboratory of Bedrock Geology, Nature Research Centre, Akademijos str. 2, LT-08412 Vilnius, Lithuania

(Manuscript received September 12, 2018; accepted in revised form March 25, 2019)

Abstract: The mid–late Homerian Age of the Silurian Period was a time of intense changes in biota, oceanic chemistry,

and sea level and is known as the lundgreni extinction (for the graptolite extinctions), the Mulde bioevent (for the conodont

turnover event) or the Homerian carbon isotope excursion (CIE) probably related to glacially influenced climate

perturbation. New information on this interval from the deep water sedimentary and graptolite succession of the Kielce

Region (Holy Cross Mountains, Poland) of the northern margin of the Małopolska Block is presented here based on

analysis of the Prągowiec Ravine section. The lundgreni–nilssoni graptolite biozones interval have been recognized

there. This interval is composed by dark shales with very rare benthic fauna, which indicate the deep open-marine (pelagic)

paleoenvironment. Ten samples were taken for the δ

org

analysis from the lundgreni (2 samples), parvus (2 samples),

praedeubeli (2 samples), praedeubeli–deubeli (1 sample), ludensis (2 samples) and nilssoni (1 sample) biozones. According

to the δ

org

results, the first positive δ

org

excursion of the Mulde Bioevent is well recognized. The δ

org

values rise

from −30.7 – −30.1 ‰ in the lundgreni Biozone to −29.3 – −28.7 ‰ in the parvus Biozone and fall below −30 ‰ in

the praedeubeli–deubeli interval. The second positive δ

org

peak of the Mulde Event was not recognized in the Prągowiec

Ravine. Based on the numerical comparisons using Raup-Crick metric of co-occurrences of graptolite species, the upper

Homerian was characterized by significant between-biozone turnover of these taxa at the given locality.

Keywords: Poland, Holy Cross Mountains, Silurian, Mulde Event, geochemistry, δ

org

Introduction

The Silurian was one of the most unstable periods in the Paleo-

zoic, marked by significant environmental changes and biotic

perturbations (Crampton et al. 2016). One such episode is in

the late Wenlock Epoch (mid–late Homerian). Graptolite

workers refer to the onset of this episode as the lundgreni

extinction (Koren’ 1987) or the “Große Krise” (Jaeger 1991),

and conodont workers refer to the immediately preceding

changes in conodont assemblages as the Mulde Event (e.g.,

Jeppsson et al. 1995) or Homerian carbon isotope excursion

(CIE; Calner 2008). Although it is debatable whether the Mulde

Event had a significant impact on conodont communities (e.g.,

Radzevičius et al. 2014c; Jarochowska et al. 2018), it has been

determined that, at least in Baltica, the community structure

and abundance fluctuation patterns radically changed as a con-

sequence of the events (Spiridonov 2017; Spiridonov et al.

2017a). The impact on the microphytoplankton communities

of the Mulde Event is not so clear and the changes in their

taxonomic composition and the size distribution probably

were more related to sea level changes (Porębska et al. 2004;

Venckutė-Aleksienė et al. 2016; Spiridonov et al. 2017b).

A twin-peaked positive carbon isotope excursion has been

documented globally in the upper Wenlock. Such carbon

isotope excursions of this age are documented: in Baltica

(Samtleben et al. 1996; Wenzel & Joachimski 1996; Bickert et

al. 1997; Kaljo et al. 1997, 1998, 2007; Samtleben et al. 2000;

Porębska et al. 2004; Martma et al. 2005; Calner et al. 2006b,

2012; Jarochowska et al. 2014, 2016a, 2016b; Radzevičius et

al. 2014c, 2016; Jarochowska & Munnecke 2016; Makhnach

et al. 2018); Laurentia (Saltzman 2001; Noble et al. 2005;

Cramer et al. 2006; Lenz et al. 2006; Sullivan et al. 2016);

Avalonia (Corfield et al. 1992; Marshall et al. 2012; Blain et

al. 2016; Fry et al. 2017); Timan (Shebolkin & Männik 2014);

Perunica (Frýda & Frýdová 2014, 2016); and Gondwana

(Vecoli et al. 2009). Thus, most of the stable carbon isotopic

data are from the Baltica, Laurentia and Avalonia paleoconti-

nents.

The purpose of the investigation is to document biostrati-

graphy, constrain the stable carbon isotopic trends of the mid-

dle–upper Homerian, and to enlighten the patterns of graptolite

community change in the Kielce Region of the Holy Cross

Mountains, Poland. Here we present the first stable carbon

isotope (δ

org

) data linked to a biostratigraphical framework

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, 2019, 70, 3, 209–221

for the Kielce Region (the Małopolska Block), and the Home-

rian biogeochemical event in the deep water open marine

facies environments.

Geological background

The Holy Cross Mountains (HCM) are located in central

Poland (Fig. 1A) and expose Paleozoic rocks in the central

part of the Teisseyre–Tornquist Zone (TTZ) which extends

from the North Sea in the NW to the Black Sea in the SE.

According to the differences in the stratigraphic, lithological,

facies, and tectonic evolution of the Lower Paleozoic, the HCM

are divided into the Łysogóry Region or Unit in the north

(Fig. 1B) and the Kielce Region (part of Małopolska Block)

in the south (Fig. 1B) (Dadlez et al. 1994).

The Łysogóry Unit is considered to be a passive margin of

Baltica (Dadlez et al. 1994; Narkiewicz 2002). The Silurian

succession of the Łysogóry Unit is without significant gaps

(Modliński & Szymański 2001) and comprises lower Llando-

very deep open-marine (pelagic) to uppermost Přídolí lagoo-

nal/fluvial (continental) deposits (Kowalczewski et al. 1998;

Kozłowski 2003). Most natural outcrops of Silurian strata are

exposed in the “Silurian Zone” (Fig. 1B) of the Łysogóry Unit

(Kozłowski 2008). The upper Ludlow positive carbon isotope

excursion associated with the Lau Event was first documented

from the “Silurian Zone” (Fig. 1B) of the Łysogóry Unit by

Kozłowski & Munnecke (2010) but no data from the upper

Wenlock were presented.

The Kielce Region is on the northern margin of the Mało-

polska Block (Fig. 1A). There are different opinions regarding

the development of the Małopolska Block: (1) it has a peri-

Gondwanan origin, rifted in the Cambrian and amalgamated

with the margin of SW Baltica during the Cambrian–Early

Ordovician (Bełka et al. 2002; Walczak & Belka 2017); and

(2) the Małopolska Block originated near the present SW

margin of Baltica (Cocks 2002; Nawrocki et al. 2007).

However, the Kielce and the Łysogóry regions were paleogeo-

graphically separate sub-basins in the Silurian (Kozłowski

2008). The Łyso góry Region was located relatively distally

and the Kielce Region more proximally in relation to the same

orogen, in other words the Łysogóry Region was closer to

Baltica then the Kielce Region (Kozłowski et al. 2004, 2014).

The Silurian sequence in the Kielce Region has numerous

stratigraphic gaps and is represented by lowermost Llandovery

deep open-marine deposits through to upper Ludlow turbi-

dites (Kozłowski & Tomczykowa 1999). Perhaps because of

the domi nance of clastic sedimentation, stable carbon isotopes

have not been investigated in the Kielce Region.

The exposed upper Wenlock and Ludlow succession is about

600 m thick (Tomczyk 1962) and is divided into the upper

Bardo, Prągowiec, and Niewachlów beds (Fig. 1C). The upper

Bardo beds are dark yellow and brown clayey shales belon-

ging to the Cyrtograptus lundgreni and probably Pristiograptus

parvus (see below) graptolite biozones. The Prągowiec beds

are composed of dark grey silty shale with rare limestone

concretions and abundant graptolites from the Gothograptus

nassa to Saetograptus leintwardinensis biozones (Tomczyk

1962). The Niewachlów beds are composed of medium-

grained greywackes with mudstone interbeds (Malec 2001)

with Bohemograptus bohemicus (Barrande), B. bohemicus

tenuis (Bouček) (Tomczyk 1962) and trilobites of Ludfordian

age (Tomczykowa 1993). The Bardo Diabase occurs between

the Prągowiec and Niewachlów beds. Using

Ar–

Ar isotope

dating its age is either 424 ± 6 Ma – 415 ± 2 Ma latest Ludlow

and earliest Lochkovian (Nawrocki et al. 2013). The igneous

intrusion is spatially separated from the sampled outcrops.

Good preservation of organic skeletons of graptolites points to

the absence of significant contact metamorphism in the stu-

died part of the section.

Material and methods

About 50 samples from the upper Wenlock and lower

Ludlow (lundgreni–nilssoni biozones) were collected from

the Prągowiec Ravine (50°44’46.04” N, 21°01’46.77” E)

located in the Kielce region (Małopolska Block) of the HCM

(Fig. 1B) about 2 km north of Bardo village on the northern

limb of the Bardo Syncline (Kozłowski et al. 2017).

Material for geochemical analysis has been selected from

five small outcrops in the Prągowiec Ravine (Fig. 1C).

The height of outcrops varies from 0.5 to 2 m. Ten samples

with well-preserved graptolites were selected from the upper

Bardo and the Prągowiec shales for stable carbon isotope

analysis. Graptolites from these shales are important for

precise biostratigraphy. Sampling at high-resolution (e.g., each

0.1 to 1 m) was not possible because the ravine is overgrown

and is covered by recent mudflow deposits and anthropogenic

debris and at the present is far away from H. Tomczyk’s (1962)

interpretation (Fig. 1C). Due to these factors and also to

the complex folding and faulting of the shales, the exact

superposition of layers below biozonal level is impossible

to determine. Due to these severe constraints on the availa-

bility of the material, the resolution of carbon isotopic sam-

pling in this study is kept approximately at two samples

per biozone (see below), where biozones were determined

based on abundant graptolite material. All the material is

stored in the Geological Museum of Vilnius University,

Lithuania.

Stable carbon isotope (δ

org

)

Samples are mostly composed of terrigenous (shale) mate-

rial with different carbonate content. For the purpose of

the δ

org

analysis, the samples that were powdered were

the same as those from which fossils were identified (total

10 samples). Approximately 0.9 mg of sample powder was

used from each sample. Powder was dissolved using 5 N

(mass equivalents) HCl acid for 24 hours at the room tempera-

ture to remove carbonate minerals. After that, the powder

residue was washed with distilled water and dried.

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GRAPTOLITE IN THE SILURIAN SHALES OF THE HOLY CROSS MOUNTAINS (POLAND)

GEOLOGICA CARPATHICA

, 2019, 70, 3, 209–221

After the pre-treatment removal of carbonates, all samples

(at least two per sediment sample) were weighed and wrapped

in tin capsules. The prepared samples were combusted with

combustion module (Costech Analytical Technologies, Inc.)

connected via Picarro Liaison Interface A0301 to the laser-

based Picarro Cavity Ring-Down Spectrometer G2121-i.

The stable carbon isotope ratio was measured in CO

and pre-

sented as per mil deviations from internationally accepted

stan dards with the reproducibility of ±0.3‰ for δ

C. The inter-

national standards (IAEA-600, IAEA C1, IAEAC2 and SRM

4990C) were used for the calibration of the reference gas

(CO

Multivariate comparison of graptolite assemblages

In order to compare the compositional changes in graptolite

assemblages in the Prągowiec Ravine, two methods were

employed: non-metric multidimensional scaling and the com-

parison of turnover within the zones and among them. We

employed a non-metric multidimensional scaling technique,

a multivariate dimension reduction technique which is robust

in revealing non-linear gradients in species composition

(Patzkowsky & Holland 2012). We used the PAST program

and the Raup-Crick metric (Hammer & Harper 2008), which

is suitable for comparison of samples with differing and

unknown abundances (Raup & Crick 1979). In our case,

the latter property is especially desirable, since the abundance

of graptolites is very difficult to measure, especially if there

are variations in preservation (e.g., many fragmented rhab-

dosomes) and in the sizes of rock samples. In order to test

the level of between-biozonal turnovers to inside-biozonal

turnovers of graptolites in the Prągowiec Ravine, we per-

formed all possible pair-wise comparisons between samples in

a given biozone c = (N

− N

)/2, where N

is the number of

samples in the first biozone) and between all the samples

between two biozones c = (N

× N

), where N

is the number of

samples in the second biozone). For this purpose, as in the pre-

vious case, we used Raup-Crick compositional distance, which

Fig. 1. A — Simplified structural map of Central Europe (Bełka et al. 2002). HCM — The Holy Cross Mountains; MGCH — Mid German

Crystalline High; OZ — Odra Zone; TBT — Tepla–Barrandian Terrane; USM — Upper Silesian Massif. B — Distribution of Silurian rocks in

the Holy Cross Mountains area (Kozłowski et al. 2014) and the Prągowiec Ravine location. C — stratigraphical interpretation of the Silurian

succession in the Prągowiec Ravine (Tomczyk 1962, fig. 9). D — Possible correlation of the Prągowiec Ravine Upper Wenlock graptolite

biozones (Tomczyk 1962) with those of revised Lithuanian (Radzevičius 2006).

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was calculated in the Vegan package (Oksanen et al. 2018) for

the R programming environment (R Development Core Team

2015). Later on, the yielded distributions of compositional

distances were compared with each other using the non-para-

metric Mann-Whitney test.

The analysis was performed on the graptolite occurrence

data from the praedeubeli, deubeli and ludensis biozones

(spanning most of the upper Homerian), since only those

biozones had more than two collected samples each and

were represented by sufficiently abundant graptolite material.

The praedeubeli Biozone was represented by 15 samples,

the deubeli Biozone by 12 samples, and the ludensis Biozone

by 18 unambiguously assigned samples with abundant grapto-

lite material. The single occurrence of Semigothograptus cf.

meganassa was not used in these multivariate analyses because

of the ambiguity of its species assignment.

Graptolite biozones

H. Tomczyk (1962) distinguished the graptolite biozones in

the Prągowiec Ravine (Fig. 1C), describing six graptolite

biozones in the Homerian–lower Gorstian interval (Fig. 1D).

Some of these biozones are not used at present. The Prągowiec

Ravine graptolite revision and new graptolite biozones cor-

relation of Tomczyk’s biozones were given by E. Porębska (in

Masiak 2010, fig. 20). At the same time as the above biozo-

nations were being developed, S. Radzevičius (2006) distin-

guished graptolite assemblages and recommended the use of

seven graptolite biozones (Fig. 1D) in the lundgreni–nilssoni

interval of the Prągowiec ravine.

Cyrtograptus lundgreni Tullberg (Fig. 2A), Monograptus

flemingii (Salter) (Fig. 2B); Monoclimacis flumendosae

(Gortani) (Fig. 2C) and Pristiograptus pseudodubius Bouček

occur in the Bardo beds (sample VU-U-10). M. flemingii,

Mcl. flumendosae and P. pseudodubius are long ranging

species, which appeared in the middle Sheinwoodian above

the riccartonensis Biozone (Zalasiewicz et al. 2009) and

disappeared in the middle Homerian during the lundgreni

Event (Koren’ 1987). C. lundgreni is the index species and its

range defines the lundgreni Biozone (Zalasiewicz & Williams

1999).

The only Testograptus testis (Barrande) (Fig. 2D) identified

was in the Bardo bed sample VU-U-6. A testis Biozone is

recognized between the lundgreni and nassa biozones in

the HCM (Tomczyk 1958; Ryka & Tomczyk 1959) as well in

the Prągowiec ravine (Tomzcyk 1962). T. testis appeared later

than Cyr. lundgreni, which accompanies T. testis until the lund

greni extinction. Therefore, the interval with T. testis repre-

sents the upper part of the lundgreni Biozone (Zalasiewicz et

al. 2009) and is sometimes referred to it as the upper subzone

(Jaeger 1991; Štorch 1994).

Pristiograptus parvus Ulst (Fig. 2E) and fragments of

Gothograptus nassa (Holm) (Fig. 2F) occur in sample VU-U-7

which is composed of dark yellow clayey shales, probably of

the Bardo beds. G. nassa ranges from the lower part of parvus

Biozone to the middle part of the praedeubeli Biozone

(Kozłowska et al. 2009) or the parvus–nassa interval (Maletz

2010). The short-ranging P. parvus appears after the lundgreni

extinction and defines the parvus range Biozone (Ulst 1974).

Gothograptus nassa (Fig. 2H) and a trilobite pygidium of

Odontopleura cf. ovata Emmrich (Fig. 2G) occur in Bardo

beds sample VU-U-9. G. nassa ranges through the parvus–

praedeubeli interval. A mass occurrence of benthic fauna

(e.g., Odontopleura) marks the lower part of the nassa Biozone

in the Prągowiec ravine (Tomczykowa 1957) and Bartoszyce

IG 1 borehole (Porębska et al. 2004) in Poland. According to

Calner et al.’s (2006a) data, O. ovata was recovered from just

above the Grötlingbo Bentonite on Gotland. The Grötlingbo

Bentonite is widespread in Laurussia at a stratigraphic level

corresponding to the parvus Biozone (Kiipli et al. 2008).

Colonograptus praedeubeli (Jaeger) (Fig. 2K) and G. nassa

(Fig. 2L) occur in the Prągowiec beds (sample VU-U-2).

C. praedeubeli ranges from the praedeubeli Biozone to the mid-

dle of the ludensis Biozone (Koren’ 1991), but is most com-

mon in the praedeubeli Biozone. Col. praedeubeli (Fig. 2M)

also occurs in sample VU-U-8 (Prągowiec beds).

Colonograptus praedeubeli (Fig. 2O) occurs in sample

VU-U-5 (Prągowiec beds). There is Col. cf. deubeli (Jaeger)

(Fig. 2N) identified in the same sample. Col. deubeli is charac-

terized by funnel- or trumpet-like sicula with distinct dorsal

process and rapid increase in rhabdosome width (Koren’ &

Suyarkova 1994). Our Col. cf. deubeli specimen has a mode-

rately expanded sicular aperture, but the width of rhabdosome

increases gradually. Col. cf. deubeli is similar in terms of rhab-

dosome width to Pristiograptus idoneus Koren’, but differs in

the form of the sicula. The sicula of P. idoneus is strongly ven-

trally curved (Koren’ 1992). Some fragments of retiolitids

with the nassa type of apertural hoods have been found in

the same sample (Fig. 2P). There is G. nassa, Semigothograptus

meganassa (Rickards & Palmer 2002) and Neogothograptus

eximinassa Maletz with genicular hoods of the nassa type in

the parvus–ludensis interval (Kozłowska 2016). The flattened

and poor preservation specimens complicate identification.

Retiolitids are distinguished based mostly on isolated, 3D

material (Lenz & Kozłowska 2007). However, N. eximinassa

marks the ludensis Biozone (Maletz 2008), S. meganassa and

G. nassa have similar stratigraphic ranges, and are confined

to the parvus–deubeli interval (Kozłowska et al. 2009;

Kozłowska 2016). The main difference that can be distin-

guished in the poorly preserved material between S.

meganassa

and G. nassa is in the width of rhabdosome. S. meganassa is

wider than G. nassa (Kozłowska-Dawidziuk et al. 2001;

Rickards & Palmer 2002) and N. eximinassa as well. Our speci-

men is relatively wide (approximately 2 mm) but the struc ture

of apertural hoods is invisible and the proximal end of rhabdo-

some is absent. Based on this, this specimen has been identi-

fied as Semigothograptus cf. meganassa. However, the index

species Col. deubeli, the first appearance of which marks

the base of the deubeli and ranges to the middle of the ludensis

Biozone (Jaeger 1991; Koren’ 1991), is not found in the sam-

ple. Therefore, the graptolite assemblage of VU-U-5 sample is

interpreted as belonging to the praedeubeli–deubeli interval.

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P. frequens appears in the nassa Biozone and disappears in

the leintwardinensis Biozone (Urbanek et al. 2012) or above

the tenuis Biozone (Štorch et al. 2014). Col. gerhardi appears

in the upper part of the ludensis Biozone and disappears in

the lower part of the nilssoni Biozone (Kozłowska-Dawidziuk

et al. 2001; Radzevičius & Paškevičius 2005) but dominated

in the upper part of the ludensis Biozone (Štorch et al. 2016).

An approximately 10 cm long mesial rhabdosome fragment

of Neodiversograptus nilssoni (Barrande) (Fig. 2R) was found

in the sample VU-U-4 (Prągowiec beds). N. nilssoni indicates

the lower part of the Ludlow and is the index species for

the nilssoni Biozone (Zalasiewicz et al. 2009).

In summary, the samples for the δ

org

analyses come from

lundgreni, parvus, praedeubeli–deubeli, ludensis and nilssoni

biozones (Table 1). There are no findings of typical Col. Deu

beli, the index species of the deubeli Biozone, in the studied

isotopic samples. Graptolites described in the samples VU-U-2,

VU-U-8 and VUU-5 have long range and can be discovering

in both praedeubeli, deubeli biozones. According to that,

the praedeubeli–deubeli interval is not split into the separate

biozones there.

Results and discussion

Turnover in graptolite assemblages

Although some indications of the community turnover can

be distinguished from the range charts of the graptolite spe-

cies, the ecological significance of those changes is not espe-

cially obvious. The composition of ecological communities

is expressed as the identity of species and their abundance.

Therefore, even if we observe a very similar set of species in

two consecutive zones, if the relative frequencies of species in

these zones are very different the final compositions in these

two zones will be very ecologically different. As a result,

numerical comparisons of sets of assemblages between zones

can be used in statistical testing of significance of temporal

changes. If the turnover between zones is continuous (when

compositions change continuously without breaks), we should

expect the fossil assemblages to significantly overlap in

the multivariate compositional spaces. On the other hand if

there are sharp changes in assemblage composition between

zones we should expect clear separation.

The comparison of distances between samples in the prae

deubeli Biozone Median = 0.21, SD = 0.21 with distances bet-

ween samples of praedeubeli and deubeli biozones

Median = 0.658, SD = 0.21 revealed their highly significant

difference, p < 0.01 (Fig. 3A). Similarly, there is a highly sig-

nificant difference between sample distances in the deubeli

Biozone Median = 0.211, SD = 0.16 and between distribution

of distances between samples from the deubeli and ludensis

biozones Median = 0.817, SD = 0.11, p < 0.01 (Fig. 3B). A simi-

lar pattern of divergence can be seen between assemblage dis-

tances in the ludensis Biozone Median = 0.249, SD = 0.27, and

the assemblage compositional distances between deubeli and

ludensis biozones, Median = 0.817, SD = 0.11, p < 0.01 (Fig. 3C).

Although an overlap in distances can be observed between

consecutive biozones, there is strong separation between

assemblages. Moreover, compositional distances between bio-

zones show strong modality, which points to the conclusion

that biozonal boundaries represent genuine turnover episodes

in development of graptolite faunas in the area represented by

samples from the Prągowiec Ravine. Additionally, it appears

that the differences in graptolite assemblages were on average

greater between the deubeli and ludensis biozones (difference

between medians of distances = 0.448) than between the prae

deubeli and deubeli biozones (difference between medians of

distances = 0.547).

The same conclusion of between-zonal differentiations in

graptolite assemblages can be drawn based on the results of

the non-metric multidimensional scaling analysis of the com-

position of graptolite assemblages (Fig. 4). The assemblages

from all three analysed biozones are very clearly distinguished,

with graptolite communities from the praedeubeli and deubeli

biozones being more closely related to each other, as was sug-

gested by the results of the previously presented pair-wise

comparative analyses. Based on the taxonomic compilation of

Sample No.

Sampling sites GPS coordinates

Formation

Biozone

org

Fauna

VU-U-10

A00

50°44’46.2” N, 21°02’16.0” E

Bardo beds

lundgreni

−30.7

M. flemingii, Mcl. flumendosae,

P. pseudodubius, Cyr. lundgreni

VU-U-6

A00

50°44’46.2” N, 21°02’16.0” E

Bardo beds

lundgreni

−30.1

T. testis

VU-U-7

A00

50°44’46.2” N, 21°02’16.0” E

Bardo beds

parvus

−29.3

P. parvus, G. nassa

VU-U-9

A00

50°44’46.2” N, 21°02’16.0” E

Bardo beds

parvus

−28.7

G. nassa, Odontopleura cf. ovata

VU-U-2

A08

50°44’46.8” N, 21°02’10.8” E

Prągowiec beds

praedeubeli

−30.2

Col. praedeubeli, G. nassa

VU-U-8

A04

50°44’48.7” N, 21°02’01.9” E

Prągowiec beds

praedeubeli

−30.8

Col. praedeubeli

VU-U-5

A07

50°44’47.6” N, 21°02’08.1” E

Prągowiec beds

praedeubeli–deubeli

−30.4

Col. praedeubeli, Col. cf. deubeli,

S. cf. meganassa

VU-U-3

A04

50°44’48.7” N, 21°02’01.9” E

Prągowiec beds

ludensis

−31.2

P. virbalensis, Col. ludensis

VU-U-1

A06

50°44’49.3” N, 21°01’58.1” E

Prągowiec beds

ludensis

−30.2

P. frequens, Col. gerhardi

10 VU-U-4

A06

50°44’49.3” N, 21°01’58.1” E

Prągowiec beds

nilssoni

−30.5

N. nilssoni

Table 1: Isotopic data for the Prągowiec Ravine samples.

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Koren’ (1991), the ludensis Biozone witnessed significant

restructuring in diversity of graptolites–retiolitids increased in

species richness and monograptids a decline in species rich-

ness. Similarly, in the Canadian sections described by Lenz

(1994, 1995), there was a significant turnover of monograp-

tids between the deubeli and ludensis biozones. According to

the global sequencing of the graptoloid clade, the ludensis

Biozone marked an interval of significant increase in grapto-

lite diversity which ended in the early Ludlow (Cooper et al.

2014). At least two large extinction events separated by hun-

dreds of thousands of years occurred in the late Homerian, not

including the lundgreni extinction event (Crampton et al.

2016). Thus the intensive graptolite turnover described in

the Prągowiec Ravine is part of the wider pattern.

Trend in δ

org

values

The δ

org

values are low and vary from −31.2 ‰ to

−28.7 ‰; (Table 1). Similar variations in upper Homerian

org

values are recorded from the part of Poland correspon-

ding to the West of Baltica (Porębska et al. 2004; Sullivan et

al. 2018). The highest variation of δ

org

values is found in

the upper Bardo beds. The values rise from −30.7 ‰ and

−30.1 ‰ in the lundgreni Biozone to −29.3–28.7 ‰ in the

parvus Biozone (Fig. 5) and drop in the Prągowiec beds and

fluctuate overall by 1 ‰ between −31.2 ‰ and −30.2 ‰

through the praedeubeli – nilssoni Biozone interval (Fig. 5).

There are two positive δ

carb

peaks in the mid-upper

Homerian (Cramer et al. 2011; Melchin et al. 2012). The first

carb

peak is found in the parvus–nassa biozones and second

one in the ludensis Biozone. Such interpretation is based on

material from the West Midlands (England) and Gotland

(Cramer et al. 2012). The West Midlands sections are domi-

nated by shallow marine facies with very rare graptolites and

the correlation of these localities is based only on conodont

biostratigraphy, sequence stratigraphy and high-precision zir-

con (U–Pb) dating of bentonites. Therefore, bearing in mind

all the possible uncertainties in correlation, the incorporation

of δ

carb

excursion with graptolite biozones should be impre-

cise. In the material from the Viduklė–61 borehole (Lithuania)

with a continuous graptolite sequence, the first positive excur-

sion represents the parvus Biozone to lower praedeubeli

Biozone interval and the second positive excursion represents

the deubeli Biozone to lower ludensis Biozone (Radzevičius et

al. 2014a) for the Gėluva Regional Stage in Lithuania (Fig. 5).

After graptolite revision from the West Midlands, the same

conclusions have been reached by Fry et al (2017).

In the gamma log data from the Viduklė–61 well, two cycles

of ~ 16.7 m and five with ~ 6.7 m period lengths were deter-

mined in the Gėluva interval (Fig. 5) (Radzevičius et al.

2014b). Similar cyclicities were found in other mid-upper

Homerian geological sections of West Lithuania (Radzevičius

et al. 2017). The Gėluva age corresponds to the mid-later

Homerian, namely from 428.45 ± 0.35 Ma to 427.86 ± 0.32 Ma

(total ~ 0.59 Ma) (Cramer et al. 2015). Based on these dates,

long cycles are about 0.3 Ma and short about 0.12 Ma.

The duration of cycles alone could be tentatively interpreted

as the 4

and 5

order cycles which are close to the Milan-

kovitch eccentricity cycles generated by the orbital forcing

(Miall 2010).

Two stage slices (Ho2 and Ho3) have been distinguished in

the mid-upper Homerian (Cramer et al. 2011) or the Gėluva

Fig. 3. Raup-Crick distances of graptolite assemblages from the Prągowiec Ravine (p-values show probabilities of null hypothesis for equality

of medians): A — between samples of the praedeubeli Biozone (c =105), and between samples of the praedeubeli and deubeli (c =180) biozones

(p < 0.01); B — between samples of the deubeli Biozone (c = 66), and between samples of the deubeli and ludensis (c = 216) biozones (p < 0.01);

C — between samples of the ludensis Biozone (c =153), and between samples of the deubeli and ludensis (c = 216) biozones (p < 0.01). Arrows

point to the medians for each distribution of between sample distances. Here c = the number of pairwise comparisons in each category.

Fig. 4. Non-metric multidimensional scaling of graptolite assem-

blages from the Prągowiec Ravine (Stress = 0.26).

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, 2019, 70, 3, 209–221

Regional Stage in Lithuania. A stage slice is an informal strati-

graphic unit that is defined on the basis of biochemostrati-

graphy. According to Cramer et al. (2011), the base of the Ho2

slice lies within the parvus Biozone and includes the first peak

of the Mulde δ

C excursion. The Ho3 slice ranges from

the base of the ludensis Biozone to the base of the nilssoni

Biozone with the second peak of the Mulde δ

C excursion.

P. parvus are found from the upper Bardo Beds in the Prągowiec

Ravine. According to that, the lower Ho2 slice boundary

doesn’t directly corresponding to the Bardo and Prągowiec

beds boundary. In the cyclostratigraphic data from West

Lithuania (Radzevičius et al. 2017), the boundary between

Ho2 and Ho3 is coincident with the boundary between Gėluva

4,1 and Gėluva 4,2 (4

order) sedimentary cycles.

It has previously been hypothesized that the evolutionary

turnover of conodonts was spread through the upper Homerian

(Jeppsson et al. 1995), although more detailed integrated ana-

lysis of interregional data revealed that the first appearances

of zonal taxa, O. bohemica longa, K. ortus absidata and

Ctenognathodus murchisoni converge to the beginning of

the Mulde event as indicated by the integrated stratigraphy

(Radzevičius et al. 2016). Therefore appearances of those taxa

in different regions should be highly diachronous (see in fig. 5

“delayed dispersal phase model”, which shows an initial phase

of evolutionary changes which is followed by the longer dis-

persal phase of conodont taxa). On the other hand, based on

the analyses of local communities, it was shown that there was

no significant community turnover in conodonts at the pro-

posed boundary of the Mulde event (Jarochowska et al. 2018).

Since there is positive evidence on evolutionary taxic disap-

pearance and appearance (Jeppsson et al. 1995; Radzevičius et

al. 2016), this lack of congruence could be explained by dis-

proportionate effects of the Mulde turnover event on rare taxa.

On the other hand, although the composition was similar in

conodonts before and after the Mulde event, their communi-

ties changed to a more even and simple abundance distribu-

tion, and there was a shift to lower abundance and higher

autocorrelation of abundance fluctuations which points to

the transition to differing community states in the upper

Homerian in the conodont clade (Spiridonov 2017; Spiridonov

et al. 2017a).

On the other hand, range data for graptolites (including

those presented here) and the multivariate analyses of their

cooccurrences point to the possibility that the late Homerian

Fig. 5. Mid–upper Homerian global stratigraphical scale (Melchin et al. 2012); graptolite biozones (Koren’ et al. 1996); stage slices (Cramer et

al. 2011); regional stages of the East Baltic (Paškevičius et al. 1994); Conodont turnover stages (Radzevičius et al. 2016); cyclostratigraphy in

the Viduklė-61 borehole and situation of the Ančia Member (Radzevičius et al. 2017); generalized δ

carb

curve (Cramer et al. 2011); lithology

(Tomczyk 1962); δ

org

data (this paper) and generalized distribution of graptolites (Radzevičius 2006) of the Prągowiec Ravine (HCM,

Poland).

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, 2019, 70, 3, 209–221

experienced several significant turnover events between bio-

zones. Those are possibly related to the periodic sea level per-

turbations associated with the 4

and 5

order cycles which

were clearly distinguished in the Baltic data (Fig. 5, and also

Radzevičius et al. 2017) and briefly described in this paper.

The same physical changes in environment are highly con-

gruent with Cramer et al.’s (2011) proposed stage slices of

the Homerian which are conceptually similar to assemblage

biozones (Fig. 5). Moreover, recent cyclostratigraphic–spec-

tral analytical studies revealed that external forcing due to

Milankovitch forcing was a significant factor in driving

changes in community compositions, and abundance of cono-

donts (Spiridonov et al. 2016, 2017a), as well as macroevolu-

tionary diversity of graptolites during the Ordo vician and

Silurian (Crampton et al. 2018). The confluence of evidence

points toward the dominance of the “common geological

cause” (sensu Peters & Foote 2002) mechanisms of change in

stratigraphy, ecology and macroevolution of biota in the

Silurian.

The first δ

org

excursion is well represented in the parvus

Biozone in the Prągowiec Ravine and related to the trilobite

mass occurrence interval in the upper part of the Bardo beds

(Fig. 5). This interval could be correlated with upper part of

microlaminated, varve-like marlstone of the Ančia Member in

the Lithuania (Radzevičius et al. 2014a) and with the lower

part of Ho2. The second δ

org

excursion is very indistinct.

This could be related to the deep marine facies which domi-

nate the sedimentary record at the Prągowiec Ravine. A simi-

lar situation is found in the Zwierzyniec-1 borehole from deep

marine facies of South-East Poland (Sullivan et al. 2018).

The magnitudes of δ

C excursions are higher in shallow water

than in deeper water settings (Noble et al. 2005), which makes

it more difficult to distinguish them in offshore shales.

On the other hand, it can be related to low sampling resolution.

However, the present results are consistent with previous

observations and show that the δ

org

data can be used as

a supporting data for stratigraphic correlation even in terrige-

nous deep water facies.

Conclusions

The first positive Homerian δ

org

excursion peak is well

represented in the Prągowiec Ravine and is in the parvus

Biozone (upper part of Bardo beds). The upper Bardo beds can

be correlated with the lower part of the Ho2 stage slice and

the Ančia Member in the Baltic Silurian Basin as well.

Nevertheless, the δ

org

curve shows a broadly similar trend

to that represented in the global Homerian δ

carb

curve and

can be used for the correlation between these datasets.

Additionally, the quantitative analyses of graptolite samples

revealed that there was a significant turnover in this planktic

group between biozones in the late Homerian (post lundgreni

Event) at the studied site. The greatest turnover occurred

between the deubeli and ludensis biozones in the Prągowiec

Ravine area.

Acknowledgements: We thank colleagues J. Mažeika and

R. Paškauskas from the Nature Research Centre (Lithuania)

for undertaking isotope analyses. We sincerely thank

M. Whittingham for the English correction A.S.’s research

is supported by the Research Council of Lithuania grant

No. 09.3.3-LMT-K-712-02-0036. This research was supported

by the Open Access to research infrastructure of the Nature

Research Centre under the Lithuanian open access network

initiative. This is a contribution to “IGCP 652: Reading geo-

logical time in Paleozoic sedimentary rocks: the need for

an integrated stratigraphy” and to “Event Stratigraphy in

the Silurian Sedimentary Basin of Lithuania”, a Vilnius Uni-

versity project.

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Zone/taxon

P. ludensis

P. frequens

C. gerhardi

P. virbalensis

C. deubeli

Gothograptus sp.

P. praedeubeli

praedeubeli

deubeli

ludensis

Appendix

Graptolite data used in multivariate analysis