J. Linn.SOC.(Zool.) 47, 311, p p . 5-14
With 2 plates and 12 text-$figures
Priqit~din Great Britnin
Octobcr, 1967
Vertebrate faunas and correlation of the Ludlovian-Lower
Devonian in eastern Europe
BY D. OBRUCHEV
AND
V. KARATAJDTE-TALIMAA
Palaeontological Institute, A C O ~ I of
J ~Sciences,
!J
Lenimki Prospect 33,
N ~ s ~ oUwAS, S.R.
C6lld
Geological Institute, VrublPa.vkio ntr. I , I‘ilnius, U.S.S.R.
Errol White was the first to elaborate an extensive scheme of vertical distribution of
vertebrate remains in the Anglo-Welsh IAN er Devonian (1950).This was shown later to be
valid also in Western Europe (\Vhite, 1956; Schmitlt, 1959) and comparable with Lower
Devonian successions throughout the worltl (e g. Denison, 1956 ; Obruchev, 1958 ; Friend,
1961).To White’s papers we still turn v h ( ~ n1 e n w h to correlate any sections of that age
with the English type one. Thus it seems appropriate t o dedicate a paper in this volume
to showing how on the foundation laid don 11 by 17’hitea fnrther construction can be built
up in the vast area of eastern Europe.
The present paper is based essentially on the data obtained by the junior author during
her collecting trips to Podolia and her work on correlation of cores from numerous boreholes in the Polish-Lithuanian trough.
Many differences of opinion as to the correlation of the strata at the Silurian/Devonian
boundary originate from the fact that only the type Downtonian is composed of near-shore
sediments, and it was possible t o base the stratigraphic subdivision of the whole Downtonian-Dittonian-Breconian series on tlie changes in the composition of the vertebrate
faunas.
I n those areas, however, where marine conditions continued much longer into the Early
Devcxiian (Czechoslovakia, Poland, Potlolia, Baltic states) tlie stratigraphy of the strata
younger than the English Ludlow was based on the index groups of invertebrates. It is
only possible t o correlate different sections with the English type one from the Traquairuspis zone up, as faunas identical or srriiilar to the vertebrate fauna of that zone are
discovered in Lithuania (Tilie beds), Potlolia (Czortkhw stage),on Spitsbergen (Fraenkelryggen division), and Northern Tinian Range (Eptarma beds).
Very important for the purposes o f correlation is the Ludlovian-post-Ludlovian
succession of the Polish-Lithuanian trough. Owing to the intensive deep boring in this
area a thick sequence of Lower Palaeozoic tlcposits was brought to light. The Silurian is
here very complete, being represented by all three stages, Llandovery, Wenlock and Ludlow. It is well characterized by invcrtebratcs. including graptolites, ostracods and
brachiopods, and can be compared with t h e English, Swedish, North Baltic, Podolian and
other successions (PaBkeviEius, 1958, 1959, 1963, 1965 ; Ulst, 1964; Gailite, 1964, 1965,
1966; Rybnikova, 1966; Kaljo & Sarv, 1966)
Above the Ludlovian (zones of Pristiograptus vulgaris, Neodiversograptus nilssoni,
Lobograptus scanicus, Pristiograptus t u i n ~ r c ~amn d I’agegiai beds) there follows in the
Polish-Lithuanian trough a thick succession (’1’7 1 t o 487 m) of marine deposits distinguished
by PaBkevifiius (1958) as Minija and Jiira beds. They contain a characteristic fauna of
brachiopods and ostracods. At the base of Rlinija beds graptolites werc found: Monograptus forrnosus and Pristiograptus ultirri u s (the latter after a personal communication
by I. PaBkeviEius). The presence of these graptolites makes it possible to compare the
D. OBRUCHEVAXD V. KARATAJCTE-TALIMAA
6
Minija beds with the PFidoli beds of Czechoslovakia. Czech geologists place the latter
above the English Ludlovian, and consider them, together with the Lochkov beds, to
be a marine equivalent of the English Downtonian (Horn?, 1962; BouEek & Horn$,l964).
Thus, in the Polish-Lithuanian trough, to the Downtonian should be referred the Minija
and Jiira beds.
These beds are followed in t)he section by the red-coloured Tilic beds, containing
vertebrate remains characteristic of the Trapmiraspis zone (besides this genus, also
Corva,spis cf. kingi Woodw., Tesseraspis sp., AngZaspis sp., Poraspididae inc. gen.)
(Karatajiitk-Talimaa, 1962, 1964). It is to be noted that in the axial part of the PolishLithuanian trough the transition from the marine to the red-coloured sediments is very
gradual, and one may assume that a t least in the most submerged area of the trough the
sedimentation a t the Downtonian-Dittonian boundary occurred without noticeable
breaks. To the south, east, and north of that area the thickness of the Minija and the Jiira
beds diminishes considerably, mainly a t the expense of the upper part of the sequence,
and the lithology of the rocks is changed.
Table 1. Correlation chart
BRECONIAN
Stoniskiai beds
1
OldRed
~
I
DITTONIAN
Tilie beds
JGa beds
DOWNTONIAN
N Z I
Minija beds
__.
Pagegiai beds
Pr. tumescens zone
LUDLOVIBN
, L. scnnicus zone
1 N . nilssoni zone
i
1
Kaugatuma form.
I
j
I
I
1
1
~
Paadla form.
P. vulgaris zone
Kaarma form.
___~____
WENLOCKIAN 1
1
I
~
1
!
i
As pointed out by Gailite (1964,1965, 1966),the ostracods of the Minija and Jiira beds
are nearly related to those of the Kaugatuma and Ohesaare formations of Saaremaa
(Oesel). Until recently, the vertebrate-bearing beds of Saaremaa (the Kaarma, Paadla,
Kaugatuma, and Ohesaare formations) were referred to the Ludlovian. Kaljo & Sarv
(1966),on the evidence of the distribution of invertebrates, chiefly brachiopods and ostracods, arrived a t the conclusion that the Ohesaare formation is Downtonian in age, and
may be correlated with the Jiira beds of Lithuania and Latvia. The upper part of the
Kaugatuma formation is correlated by these authors with the Minija beds.
When dissolving the detrital limestones of the upper part of the Jiira beds in 30%
monochloracetic acid some small remains of vertebrates were found : scales of TrimeroZepis lithuanica Kar.-Tal. (Text-figs 1 to 7 ; P1. 1,figs 4 to 6), belonging, to judge by their
microstructure, to the Phlebolepididae, but rather different from the older and only genus
Phlebolepis; fragments of tesserae of Strosipherus ( = Oniscolepis) sp. ; scales of Thelodus
obliteratus Gross, Th. cuneatus Gross ; acanthodian scales and spines : Nostolepis striatu
Pander, N . gracilis Gross, Gomphodus sandelensis Pander, Climatius sp., and some scales
Lower Devonian correlation in eastern E u i o p ~
c
of doubtful appurtenance. Lower down in tlie Jiira beds only remains of anaspids and
acanthodians are met with (Karataj~te-Taliriiaa,1967, in press). The vertebrate fauna of
the upper part of the JCra beds is nearly related to that of the Ohesaare formation of
Saaremaa : Strosipherus indentatus Pander, Tolypelepis u ndulata Pander, Thelodus parvidem Ag., T . costatus Pand., T . trilobatua Hoppe, T . bico9tatus Hoppe, Nostolepipis striata
Pand., Gomphodus sandelensis Pand., Cliinatius curuatus (Pand.). Thus, the composition
of the vertebrate fauna confirms the correlation of the JClra beds and the Ohesaare
formation.
The vertebrates from the underlying Ludlovian of Saarernaa are rather well known
and for a long time served as a base when detei,lniniiig the age of vertebrate faunas of other
areas. Aaloe (1963) has shown that tlzc dolomites with Tremataspis rnammillata outcropping in the quarry at Himmistc-Kriigu do not form an upper part of the Kaarma
formation, but belong t o the Kipi member in the upper part of the Paadla formation.
Accordingly, the lists of the vertebrates o f both these formations underwent changes.
That of the lower, or Kaarma formation, colnprises. Thelodus Zuevis (Pander),T . carinatus
(Pander),T . schmidti (Pander),Trematc1.spr.s schmdti Rohon. Thyestes vermcosus Eichn .,
Saaiuinnaspis mickwitzi (Rohon), Witatupis schrencki (Pander), while in the upper. or
Paatlla formation, occur, besides the three mentloned thelodonts, also T . parvidens Ag.,
Phlebolepis elegans Pander, Tremataspix ~numinillutaPatten, T . milleri Patten, T . rohotii
Patten, Dartmuthia gemmifera Patten, 0eseknpl.s pustulata Patten, Procephlaspis
oeselensis Robertson, Saarolepis oeselensis (Robertson).Although these two faunas have
only the three thelodont species in comnion. they are rather similar in composition, consisting mainly of numerous osteostracans.
I n the overlying Kaugatuma formation, a radical replacement of vertebrate faunas
takes place, The Kaugatuma fauna is restricted to the npper part of the formation, and
contains only acanthodians (Gomphodus sandelensis, Onchus tenuistriatus Ag., 0. roemeri
Hoppe) which appear here for the first time. Higher up, in the Ohesaare formation, there
are no osteostracans a t all, only two genera of heterostracans, and thelodonts and acanthodians. This sharp change in the composition of the vertebrate faunas can be partly
explained by the changes in the conditions of life and burial. The vertebrate remains of the
Kaarma and Paadla formations occur in sediments of a semi-closed basin, temporarily
freshened due to discharging of a river (Aaloe, 1963). The vertebrate remains of the Ohemarc formation and the upper part of the Kaugatuma were buried in marginal marine
deposits. Thus i t was natural that given only the succession of vertebrate faunas during
the Late Silurian in Estonia, no stratigraphic value could be attributed t o the vertebrate
remains, and all the four formations were referred to the Ludlovian.
The discovery in the marginal marine deposits of the upper part of the Jfira beds of a
Vertebrate fauna nearly related to the Ohesaare fauna facilitates the correlation of both
these formations, as well as of the underlying oncs, in the North and South Baltic areas.
The replacement of the vertebrate faunas in tlie upper part of the Kaugatunia coincides
with the boundary between the Ludlovian and the Downtonian. Further search for
vertebrate remains in marine Liidloviari and DOIF
ntonian will help to substantiate this
boundary.
The Ohesaare formation seems to corrcspontl only partly to the Jtira beds, the equivalents of the uppermost part of the latter, well a s those of the Traquairaspis zone, having
been destroyed. This is evidenced by the erratic boulders scattered throughout the North
German plain, which supplement t o a certain extent tlie Saaremaa section. Gross (1947,
1961) discerns three basic types of bould
(1)those with Strosipherus, thelodont and
acanthodian scales, ( 2 ) those with Cephalrispis and anaspid scales, and (3) boulders
Bey 36, 37, already containing Dittoriian vertebrates. Traqimiraspis, Cephalaspis?,
Corvaspis and Anglaspis.
Boulders of the first type possibly correyond to the uppermost part of tlie Jtira beds.
I n them scales of Trimerolepis seem also to have bten found ( T h e b d wsp. ind.. Gross, 1947,
8
D. OBRUCHEVAND V. KARAATAJUTE-TALIMAA
Text figs 1 to 12. Numbers refer to the collection of the Geological Institute, Vilnius.
1 and 2. Trinaerolepsis Zithuonica Kar.-Tal., No. 5-01044 ( x 70); 1, head scale-outer view;
2, head scale-inner view.
3 and 4. Trimwokpis Zithunizica Kar.-Tal., KO. 5-01045 ( x 70); 3, body scale-outer view;
4, body scale-inner view.
5 and 6. Trimwokpis Zitl~umicaKar.-Tal., No. 5-01042 ( x 70); 5, body scale-outer view;
6, body scale-inner view.
Lower Devonian correlution in eastern Europe
9
7. Trinscrolepis Zillurniiicrc Kar.-Tal., KO. 5-01163 ( x 124) : transverse section of anterior
part, of body scale. Lithuania; S t o i i i 3 h i bore-hole, 1211 to 1217 m. Jura beds, Downtonian.
8. Trimwokpis tiirmnicu. Kar.-Tal., Yo. 5.01 162 ( x 1 1 0 ) ; transverse section of anterior part
of a scale. K.Timari, r. Velikap, outvrop 32, Dcnvntoniari.
9 and 10. Trim,erolepi.s t i m a i i i c r c . Kar.-TI':il., S t ) . 5.01035 ( x 7 0 ) ; 9, body scale-outer view;
10, body scale-inner view.
11 and 12. ~ ' T i 7 n C ? d ? p i Stiinciiriccr Kw..'L11., I i o l o t ~ - v p c ,KO. 5-01013 ( x 70); 11, body scaleouter view; 12, body-scale-inner vicia.. iX.l i n i a n , r. Veliltaya, outcrop 32. Doantonian.
pl. 20 ( I ) : 9), as may be judged from t Iic f w m of the crown and the presence of a wide
pulpar depression with a small pulpar opening in the most distal part of the scale.
The vertebrates of the second type of boultl~rgive no base for a correlation. Remains
of anaspids have been met with in the niicldle part of the JGra beds of some Latvian and
Lithuanian bore-holes.
The boulders of the third type are ol)viously equivalent to the Tilie beds, or the
Truquwiraspis zone.
The vertebrates of the strata overlying tlic Tilie beds are still insufficiently known in
10
D. OBRUCHEVAND V. KARATAJ~TE-TALIMAA
the Baltic, but the recent finds of Belgicaspis crouchi (Lank.) in the Stonigkiai beds, and
of Rhinopteraspis cornubica (McCoy) ( = dunensis Roemer) and Porolepis in the Beguvis
beds allow us to correlate these formations with the Upper Dittonian and Breconian
respectively.
I n the north of the Timan range there are two vertebrate-bearing horizons. The lower
one, overlying the Upper Llandoverian Halmer beds, can be referred to the Downtonian.
It contains a fauna identical with that of the J c r a and Ohesaare beds: Trimerolepis
timanica Kar.-Tal. (Text-figs 8 to l a ) , Strosipherus indentatus, Gompliodus sandelensis.
Nostolepis striata. Onchus roemeri, Climatius curvatus. These beds were correlated with
the Ohesaare and referred first to the Upper Ludlovian (Kossovoi & Obruchev, 196%),and
later to the Lower Ludlovian (Kossovoi & Barkhatova, 1965). A considerable break
corresponding to the whole of the Wenlockian and Ludlovian separates this horizon
from the underlying beds, and a much lesser one, from the overlying Eptarma beds with
a probable Lower Dittonian fauna : Tolypelepis timanicu Obr., Traquairaspis ( 2 ) sp.,
Timanaspis kossovoii Obr.. Bidymaspis cf. grindrodi Lank., Cephalaspis sp., Onchus cf.
verus Schmidt Two scales, one of Strosipherus, and one of Trimerolepis,seem to havebeen
redeposited from the underlying Dow ntonian horizon.
I n the Podolian sequence numerous vertebrate remains appear only from the base of the
CzortkGw stage up. A t the top of the Skala stage a single specimen of Irregulureaspis has
been found, and in the limestone intercalations of the Borszczbw stage only small Nosfolepis-like scales of acanthodians have been discovered up to the present.
Dissolving samples of limestone intercalations of the Czortk6w stage produced a large
material of small vertebrate remains. Their first appearance in the section coincides with
the lower boundary of the Czortkbw stage as established on the basis of invertebrates
(Nikiforova, 1965).At the very base of the stage a few scales of Thelodus cf. scoticus Traq.,
T h . oervigi Kar.-Tal., Apalolepis obruchevi Kar.-Tal. (Pl. 1, figs 1 to 3), and fragments of
pteraspids are found. Somewhat higher in the section ( 10 to 15 m) fragments of Traquairaspis appear. Very numerous remains occur from the middle part of the stage up.
They are usually scattered throughout the limestone intercalations, but sometimes they
form accumulations on the bedding planes, together with debris of Lingula.
The composition of the vertebrate fauna throughout the whole thickness (-200 m)
of the Czortkbw stage is very uniform. The thelodonts comprise two genera, Thelodus and
Apalolepis. Some new forms of heterostracans are found, represented by head- and bodyscales: Polymerolepis whitei Kar.-Tal. (Pl. 2 , figs 1 to 4)and Seretolepis elegans Kar.-Tal.
(Pl. 1, fig. 7 ) . Rather common are small tesserae (?) bearing a single tubercle, fragments of
plates, and scales of Traquairaspis. More rare are scales and fragments of plates of
Corvaspis.
Most numerous are fragments of pteraspids. More complete plates are met with only
N
PLATE1
Numbers refer to the collectim of the Geological Institute, Viliiius.
Figs 1 and 2. Apnlolepis obrzbchevi Kar-Td., No. 5-00341 ( x 50); 1, scala-outer view; 2, scale-inner
view. Podolia, Grbdek, Czortkbw stage, Dittonian.
Fig. 3. Apnlolepiu obruchevi Kar-Tal., N o . 5-00303 ( x 50): scale-outer view. Zaleszczyki, Czortk6w
stage, Dit,tonian.
Fig. 4. Trimerolepis lithuunica Kar-Tal., KO. 5-00428 ( x 50) ; body scale-outer view.
Figs 5 and 6. Trimerolepis Zithuairica Kar-Tal., No. 5-00429,holotype ( x 50); 5, body scale-outer view;
6, body scale-inner view. Lithuania, St,oniSkiaibore-hole, 1211 to 1217 m, JUra beds, Downtonian.
Fig. 7. Seretoiepis elegans Kar-Tal., N o . 5-00409 ( x 40); fragmentary seale-auter view. Podolia,
Zaleszczyki, ‘transition beds’, Dittonian.
Fig. 8. Traquaira,.spissp. No. 5-00406 ( x 10) : outer view of scale. Podolia, Zaleszczyki, ‘transition beds’,
Dittonian.
Fig. 9. Radotina ( ? ) sp. ind., No. 5-00399 ( x 30); outer view of tessera. Podolia, Zaleszczyki, ‘transition
beds’, Dittonian.
Lower Devonian correlation (:)L eastem Europe
11
from the middle part of the stage up, while compl
dorsal shields mit'li the orbito-rostra1
area preserved occur very rarely in the upper part. Most common are determinable remains
of Podohpis podolica (Alth) ( =P. lerichci (Zych)), less so 2ascinaspi.s heintzi (Brotzen).
Only one specimen of Belgicaspis crouchi was found. Other species, known from the
literature, have not been identified.
Poraspids are found in small fragment,l;;.as well as in more or less complet'e dorsal and
ventral shields (from the middle part of t,he stsageup) and scales. According to Balabay
(1956), they all belong to a single species, Pornspis sturi (Alth).
Identifiable remains of cephalaspids are c~t~remely
rare.
Arthrodiran remains are rat,her common only in the upper part of the st'a'ge.Fragments,
isolated plates, and more complet'e part.s of the body armour of Palaeacanthaspis vasta
Brotzen are ascertained from the locality of Dobrowlany. For the first time tesserae and
rhombic scales of a rhenanid related to Hrrcfofi'm (Pl. 1, fig. 9) were found. Peculiar small
plates, named Tyriolepis radiata Kar.-Tal. (PI.2 , figs 5,B) consist entirely of bone, and are
pierced by canals ascending from the under surface nearly to the upper. There are symmetrical more or less convex plates covered with radial row's of bony tJiibercles,as well as
asymmet>ricalones of bhe same structure. Perhaps they belong to t'he Gemuendinidae.
Acant,hodian remains, occurring already as low in t'hc section as the Borszcz6w stage, are
especially abundant in the CzortkGw. Scales and syniphysial dental plates of two types
(Nostolepisand Gomphodus),fin spines of the Onchus type, and hexagonal head plates are
present practically in all limestone intercalations.
The sediments of the CzortkGw stage present an example of a marine (in their upper
part marginal) environment,. Vertebrate. rttmains were buried mostly as isolated scales,
small t,esserae and fragment's of plates. Isolatctl or fused plates of pteraspids, poraspids,
a,nd some arthrodires, occurring spora.dically from the middle part of the stage up, are
exceptions. It is reasonable tlo assume t'hat these vertebrates could inhabit marginal
marine, as well as delt'a'iczones, and some of theni, the pt,eraspids in particula'r, were most
likely euryhaline.
Such genera as Truquairaspis a,nd ( 'ormytoisin other areas (Engla'nd, Spitsbergen,
Lithuania) are connected only with coiit ineiit.al sediments (deltaic, fluvial, lacustrine).
They could be carried by rivers int,o thc? rria,riiiedeposits of the Czortktiw stage, where
t'hey are always represent.ed by isolated scales or small fragments.
On the other hand, there occur in the Czort,liO.wsuch probably marine forms as Radot i w ( 1 ) and the presumed gemuendinid Tyriolepis. Such a mixture of fresh-water, euryhaline, and marine forms lends a great stratigraphic importance to t,he vertebrate assemblage of the Czortk6w stage.
The CzortkGw stage has been consideled Downtonian or even Ludloviari in age. However, its vertebrate fauna is unmistakably Ditt'onian, as already suggested by Denison
(1056)and Obruchev (1958).The finding of pteraspicl remains at the very base of the stage
c,onfirmsthat t'he whole st'age from t'he bast. t,o t'lie top belongs t o the Lower Dittonian.
It is rea'sonable t,o suppose that the untlerlying Borszcztiw stage, which is not separated
from the Czortkbw by any visible break, is equivalent to the Downtonian, all the more so
1'I.ATE
2
Xioiibera refer to the collectioii. of the Ocological ITistiticte, Vihira.s.
Figs 1 and 2. P o l y n m o l e p i s whitei Kar-Tal., No. 5.00363 ( x 50); 1, body scale-outer view; 2, body
scale-inner view. Podolia, Dobrowlsny, ' fransition beds', Dittonian.
Fig. 3. Polymerokpis whitei Kar-Tal., No. 5.00374 ( x 5 0 ) ; head scale-outer view. Podolia, Dobromlany, 'transition beds', Dit,tonian.
Fig. 4. Polymerolepis white?: Kar-Tal., No. 5.003 I6 ( x 50): body scale-inner view. Podolia, Grbdek,
middle part of Czortk6w &age, Dittonian.
Figs 8 and 6. Tyriolepis rndiutn Kar-Tal., No. 5-00352 ( x 20); 5 , plate-out,er view; 6, plate-inner
view. Podolia, Dobrowlany, 'transition beds ', Uittonian.
(Frrci,rg 1’. 1 1 )
Plate 2
Chief localities (Dniester section)
The1odu.s cf. scoticus
TA. oerviyi
Apalolepis obruchevi
Thelodont scales
Polymerolepis whitei
Seretolepis eleyans
Traquairuspis sp. ind.
Weigeltuspis nltn
Weigeltaspis sp.
Corvaspis sp. ind.
Tesseraspis( ?) SIJ. ind.
Poraspis sturi
*Seretaspis zychi
*Irregulareaspis stensioi
*Bothrinspis kicieri
Poraspididao gem ind.
Porlolaspis podolicrc
PodolaTpis sp. i d
Pteraspis kncri (!)
Loricopterccspis &hi
Zascinaspis heiritzi
Z . maynippinealis
Zascinaspis sp. ind.
Pteraspis grncilis ( ? )
Mylopteraspis ( ? ) sp. intl.
Eelgicaspis crouchi
A lthaspis elongnta
Pteraspididae gen. ind.
C'ephalaspis kozlowskii
Ceplmlaspis sp. sp.
Palaeacanthaspis vasta
*Dobrowlunitr podolica
Tyriolepis raditrta
l i a d o t i m ( ? ) sp. ind.
Kujdanowiaspis rectiforrnis
*K. podolica
Kujdanowiaspis sp.
Nostolepis- and Gomplioclus-like scalcs
Onchus-like spines
Acanthodian head plates
Zasulince
Doroszowce
Table 2 .
Gr6dek
Koscielniki
~
* Literarv data
Dobrowlany
I
(lower part)
Czortk6w stage
I
~~~~~
-1
Iwanie (lower part)
Uscieczko (lower part)
~~
'Transition beds'
Vertical range of Dittonian vertebrates in Podolia
(upper part)
Iwanie (upper part)
Uscieczko (upper part)
.~
First zone of Old Red
Lower Deconian correlatiori in eastern Europe
13
since a Dittoniangenus, Irregulareaspi<T,
was foiintl a t the top of the still lower Skala stage.
Thus i t is evident that the so-called ‘Tiveriaii stage’, embracing both the Borszcz6w and
the Czortkhw stages, is equivalent to tlic Downtonian and the Louer Dittonian, and not
to an imaginary break between the Lutllovian and the Downtonian in the English type
succession, as supposed by Nikiforova & O h t (1961, 1983).
The vertebrate fauna of the Old Red overlying the CzortkGw stage needs a thorough
revision. The numerous species of pteraspidb and arctolepids established by previous
authors are mostly only poorly known and often iiitlistingiiishable. The last reviser of tlie
pteraspids and poraspicls, Balabay (1956, 1959, 1960, 1961) reduced the number of recognizable species (including those of the CzortkOn- stage) to 13. Balabay (1960)discerns three
zones in the Podolian Old Red (from below). (1) zone with Pteraspis lerichei, P t . major,
Bra( hypteraspis heintzi and Cephalaspis lyelli ; ( 2 )zone with Pt. major aiid Pt. elongata, and
(3) zone u itli Pt. major, Pt. elongata, Pt longirostra. and Brachypteraspis latissima.
It is evident from this enumeration that tlie ptcraspid assemblages of all three zones are
neaily identical, as Pt. lerichei Zych (=Podolaspas podolica (Alth))occurs in both the first
and second zones, P t . major Zych ( = P t . kiieri Imikester), and Pt. elongata Zych ( = A l t h aspis samsonoviczi Tarlo, 1961, =A l . elonguta (Rltli, 1874)),in all three, Pt. longirostra
Zych ( = Althaspis) and Brachypteraspis lutissinru (Zych), in both the second and third
zones Only Br. heintzi Brotzen ( =Zascinaspis) scenis to be restricted to the upper part of
the C‘zortBOw stage and to the first zonc. Moreover, many species of pteraspids and of
other vertebrates are common to the Czortlrtiw stage and t o the Old Red (Table 2).
Ptwaspis elongata and Pt. loiigirosttn were referred to Rhinopteraspis by Brotzen
(1936, see also Obruchev, 1958), but as pointed out by White (1938: 97) P t . elongata is
nearer Pt. leachi White ( = Althaspis), and tlie samc seems to apply to P t . longirostra ( c f .
Brotzen, 1933: 459). Thus, there is no wason to correlate even the highest strata of the
Podolian Old Red with the Breconian or even nith the Eifelian, as was done by Zych
(1927) and sonie other authors, but only to the iipper part of the crouchi-rostrata and to
the leachi zone.
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