This article appeared in a journal published by Elsevier. The attached
copy is furnished to the author for internal non-commercial research
and education use, including for instruction at the authors institution
and sharing with colleagues.
Other uses, including reproduction and distribution, or selling or
licensing copies, or posting to personal, institutional or third party
websites are prohibited.
In most cases authors are permitted to post their version of the
article (e.g. in Word or Tex form) to their personal website or
institutional repository. Authors requiring further information
regarding Elsevier’s archiving and manuscript policies are
encouraged to visit:
http://www.elsevier.com/copyright
Author's personal copy
Available online at www.sciencedirect.com
Palaeoworld 19 (2010) 108–128
Research paper
Palaeozoogeographical connections of the Devonian vertebrate
communities of the Baltica Province. Part II. Late Devonian
O.A. Lebedev a,∗, E. Luksˇevicˇs b, G.V. Zakharenko a
a Palaeontological Institute, Russian Academy of Sciences, Profsoyuznaya St. 123, Moscow 117997, Russia
b Department of Geology, University of Latvia, Rainis Blvd 19, Riga LV-1586, Latvia
Received 23 December 2008; received in revised form 19 October 2009; accepted 9 December 2009
Available online 21 December 2009
Abstract
Late Devonian vertebrate communities within the Baltica zoogeographical Province are analysed for intra- and interprovincial connections.
Components within the category of provincial endemics are used to assign the communities to a particular zoogeographical province. Marine
and continental, presumably freshwater types of vertebrate dispersal are outlined. During the Late Devonian marine dispersal is displayed by
ptyctodonts, struniiforms, and some dipnoans, and continental dispersal by psammosteids, acanthodians, and some arthrodires.
Isolation of communities is reflected by predominance of local and provincial endemics; the majority of polydemics and cosmopolitans records
wider connections. It is suggested that proportions of endemism-cosmopolitanism significantly correlate with sea-level changes. Augmentation
of endemism may result from regressions that increased isolation of faunas. Separation from the main sea basin may have reduced competition,
resulting in radiation within limited areas.
Constant faunal interchanges are observed between Baltica and Laurentia, but connections to other provinces are sporadic. Different vertebrate
groups indicate varying dispersal capabilities. The cosmopolitans are ptyctodont, arthrodire and antiarch placoderms, chondrichthyans, acanthodi-
ans, actinopterygians, and struniiforms. Agnathans and other sarcopterygians show limited dispersal capabilities. Continental and pelagic marine
faunas may have interacted by penetration of the members of one fauna into the habitat of another one during favourable periods.
© 2009 Elsevier Ltd and Nanjing Institute of Geology and Palaeontology, CAS. All rights reserved.
Keywords: Palaeozoogeography; Devonian; Baltica Province; Endemism; Dispersal
1. Introduction
This paper continues the analysis of the vertebrate distribu-
tion within the Baltica zoogeographical Province into the Late
Devonian presented by Luksˇevicˇs et al. (2010) for the Middle
Devonian. The dispersal model during the Late Devonian in
comparison to the Middle Devonian is exemplified by various
agnathan and gnathostome vertebrate groups. The hypothesis
on the correlation between the endemic-cosmopolitan compo-
sition of the community and sea-level changes is checked. A
survey of the interprovincial connections of vertebrate commu-
nities is used as a tool for highlighting predominant dispersal
∗ Corresponding author. Tel.: +7 495 3391988; fax: +7 495 3391266.
E-mail addresses: olebed@paleo.ru (O.A. Lebedev), ervins.luksevics@lu.lv
(E. Luksˇevicˇs).
directions and clarifying of the role of various vertebrate
groups.
2. Methods and terminology
The faunal elements of Late Devonian communities are
examined at the generic level for their distribution within
zoogeographical districts, provinces, regions, and realms. For
explanation of the newly introduced terminology and for the sug-
gested Givetian-Famennian global vertebrate regionalisation,
see Lebedev and Zakharenko (2010).
Where available, sedimentological data are used to recon-
struct environments. When tracing various dispersal directions
we take into account the physico-geographical conditions
suggested by earlier authors (Tikhomirov, 1967; Savvaitova,
1977; Sorokin et al., 1981; Rodionova et al., 1995). The
most important studied territories (Main Devonian Field and
1871-174X/$ – see front matter © 2009 Elsevier Ltd and Nanjing Institute of Geology and Palaeontology, CAS. All rights reserved.
doi:10.1016/j.palwor.2009.12.003
Author's personal copy
O
.A
.L
eb
ed
ev
et
a
l.
/P
a
la
eo
wo
rld
19
(20
10
)1
08
–1
28
10
9
Table1
EarlyFrasnianvertebratecommunitiesoftheBalticaProvince.Basedupon:Krylova(1973),Vorobyevaetal.(1997),Esinetal.(2000),ValiukevicˇiusandKruchek(2000),Moloshnikov(2007,2008)andunpublished
author’sdata.
LocalendemicsProvincialendemicsQuasiendemics/didemicsPolydemicsCosmopolitansE-Cindex
Pl¸avin¸ascommunityofMDF(Latefalsiovalis-transitansZoneinterval)
Rhamphodopsis
Platycephalichthys
Haplacanthus
“Acanthodes”
Psammosteus
Plourdosteus
Asterolepis
Homalacanthus
Eusthenopteron
Conchodus
“Rhynchodus”
Grossilepis
Glyptolepis
Laccognathus
Ctenurella
“Ptyctodus”
Bothriolepis
Strunius
Holoptychius
Latvius
Rhinodipterus
Griphognathus
Moythomasia
2:2:6:4:9
TimancommunityoftheCDF(LatefalsiovalisZone)
TartuosteusHaplacanthusPsammosteus
Plourdosteus
Asterolepis
Devononchus
“Acanthodes”
Glyptolepis
Laccognathus
“Ptyctodus”
Atopacanthus
Onychodus
Holoptychius
1:1:4:3:4
TimancommunityofSouthTiman(LatefalsiovalisZone)
cf.Nostolepis
cf.Diplacanthus
Cheiracanthus
Haplacanthus
“Acanthodes”
Psammosteus
Plourdosteus
Devononchus
Holoptychius
“Rhynchodus”
Glyptolepis
Laccognathus
Holonema
Ctenurella
Bothriolepis
Atopacanthus
Rhinodipterus
Moythomasia
3:2:4:3:6
Ust’-YaregacommunityofSouthTiman(transitansZone)
cf.Cheiracanthus
cf.Rhadinacanthus
Haplacanthus
“Acanthodes”
Psammosteus
Plourdosteus
Devononchus
GlyptolepisHolonema
Eastmanosteus
Bothriolepis
Holoptychius
Moythomasia
2:2:3:1:5
Ust’-YaregacommunityofMiddleTiman(transitansZone)
HaplacanthusPsammolepis
Psammosteus
Plourdosteus
Asterolepis
Homacanthus
Glyptolepis
Laccognathus
Ctenurella
“Ptyctodus”
Bothriolepis
Holoptychius
Latvius
Rhinodipterus
Moythomasia
0:1:4:3:7
VyucheycommunityofNorthTiman(?)
Psammosteus
Devononchus
GlyptolepisBothriolepis
Holoptychius
0:0:2:1:2
StylacommunityofUkraine,DonetsBasin(?)
PsammolepisDipterusBothriolepis
Holoptychius
Rhinodipterus
0:0:1:1:3
Author's personal copy
110 O.A. Lebedev et al. / Palaeoworld 19 (2010) 108–128
Central Devonian Field) are abbreviated in the text (MDF and
CDF).
3. Dispersal of provincial endemics within the Baltica
Province
Analysis of the distribution of provincial endemics includes
the geographical positioning of their communities, brief charac-
teristics of the environments and the balance of endemics versus
didemics/quasiendemics, polydemics and cosmopolitans. The
combination of provincially endemic genera in various districts
is used as the main characteristic of the province during the time
slice under consideration.
Local endemics provide no information on faunal inter-
changes, but characterise separate districts. Provincial endemics
define the directions of the faunal interchanges between dis-
tricts and typify the entity of the province during successive
chronological periods.
3.1. Upper Devonian
3.1.1. Early Frasnian (Late falsiovalis-transitans time)
(Table 1, Fig. 1)
During the early Frasnian the Pl¸avin¸as vertebrate community
is distributed in a shallow-water sea in the central and western
parts of the Baltic syneclise (MDF). It is widely known as “Snet-
naya Gora vertebrate assemblage” (Gross, 1933, 1941, 1942;
Ivanov, 1990; Ivanov et al., 2006). The environment changed
from lagoonal in the very western part of the basin to tidal and
wave-influenced shallow-marine in the eastern part, with rare
Fig. 1. Distribution of vertebrate communities of the Baltica Province during the early Frasnian. Black squares indicate the most important vertebrate localities.
Arrows illustrate faunal interchanges between districts; abbreviations by the arrows list the main vertebrate groups participating in the dispersal. Abbreviations:
CDF—Central Devonian Field, MDF—Main Devonian Field, ACA—acanthodians, ANT—antiarchs, ART—arthrodires, CHO—chondrichthyans, DI—dipnoans,
OST—osteolepiforms, POR—porolepiforms, PSA—psammosteiforms, PTY—ptyctodonts, STR—struniiforms. Base maps for Figs. 1–3 compiled after Vinogradov
and Nalivkin (1960), Tikhomirov (1967) and Rodionova et al. (1995).
Author's personal copy
O.A. Lebedev et al. / Palaeoworld 19 (2010) 108–128 111
invertebrates, but still diverse fishes (Tovmasyan and Stinkulis,
2008). Timan fish communities of the CDF and South Timan
dwelled in shallow, brackish-water environments, but by the end
of this time the freshwater input brought more clastic materials.
The Sargay Regional Stage is normal marine, but no vertebrates
are known from this time interval (Tikhomirov, 1967; Rodionova
et al., 1995). The Ust’-Yarega communities of the South and
Middle Timan lived under the maximum marine transgression
conditions. In contrast, the Donets Styla Formation placed close
to the Ukrainian Island is completely terrigenous, and apart from
fishes yields abundant plant remains (Krylova, 1973), suggesting
onshore, possibly deltaic or flood plain environments (Table 1;
Fig. 1).
In comparison to the Givetian (Luksˇevicˇs et al., 2010),
early Frasnian communities (Table 1) show the dominance of
cosmopolitans and a significant proportion of didemics. Provin-
cialism is characterised by the assemblage composed only of
the acanthodians Haplacanthus and “Acanthodes”. This situa-
tion contrasts with the late Eifelian (Luksˇevicˇs et al., 2010), when
the provincially limited acanthodian connection is dominated by
wider didemic and polydemic dispersal.
The holonematid Megaloplax from South Timan was col-
lected in the locality exposing the deposits of the Lower Frasnian
Ust’-Yarega and Middle Frasnian Domanik Formations and its
exact origin is unknown. More, A. Ivanov expressed an opinion
(2009, pers. comm.) that the holotype specimen belongs in fact
to Holonema, thus this genus is not used in the current analysis.
The ptyctodont placoderm Rhamphodopsis, from the Eifelian-
Givetian of East Laurentia, survived into the Frasnian in the
MDF.
This predominantly cosmopolitan distribution reflects wide
dispersal of early Frasnian faunas across the globe. Prevailing
faunal connections are traced to Eastern Laurentia (see Section
4.1.1, Section 4.1.2, Fig. 1; Table 8). The basic analysis of this
fauna was made by Ivanov (1990) and Schultze and Cloutier
(1996).
3.1.2. Middle Frasnian (punctata-jamieae time interval)
(Table 2)
The geographical distribution of middle Frasnian commu-
nities is similar to the early Frasnian, except that vertebrates
reappear in Severnaya Zemlya (Middle-Late Matusevich com-
munity) (Table 2).
During punctata Zone (Dubnik time) the MDF basin
regressed, followed by a new wide Early hassi Zone (Daugava
time) transgression. Lagoonal type hypersaline basins formed in
the west, but in the east siliciclastic deposition dominated, with a
significant freshwater input from the uplands. The jamieae Zone
(Snezha time) evidenced a complete transgression–regression
cycle, producing changeable deposits of the shallow sea
(Sorokin et al., 1981). The deposits of punctata–Early hassi
Zone (Semiluki time) in the CDF are shallow-water marine with
normal salinity (Rodionova et al., 1995).
During punctata–Early hassi Zone (Domanik time) a deep-
water meridionally directed trough formed in the South Timan.
Its deeper parts show evidence of anoxia, but on the lateral
shelf parts normal marine conditions persisted, inhabited by
a rich shallow-water biota, including vertebrates. Those were
sometimes also buried in the deeper parts of the basin (House
et al., 2000). The Kraypole community of Middle Timan and
the Ust’-Bezmoshitsa community of North Timan also lived in
shallow-marine, near-shore environments, as well as the Middle-
Late Matusevich community of Severnaya Zemlya (Matukhin
and Menner, 1999; Luksˇevicˇs, 1999).
The structure of middle Frasnian communities (Table 2)
is similar to that of the early Frasnian communities in the
dominance of cosmopolitans. The generic composition remains
virtually the same as that during the early Frasnian. Provincial
endemics include only the acanthodian “Acanthodes” and the
osteolepiform Jarvikina. Their dispersal shows interconnection
of the MDF and Timan communities. The connection of both to
the CDF is not evident and may be traced only by the didemic
arthrodire Plourdosteus.
3.1.3. Late Frasnian (rhenana-linguiformis time interval)
(Table 3)
The geographical distribution of vertebrate communities dur-
ing the late Frasnian (Fig. 2) generally remains unchanged from
the middle Frasnian. The Pamu¯sˇis-Stipinai basin of the MDF
with mainly vertebrate fossils was shallow and dominated by
sedimentation of sands followed by hyperhaline conditions with
only rare occurrence of vertebrates. During linguiformis Zone
(Amula time) the shallow sea basin existed only in a restricted
area of the western part of the MDF (Sorokin et al., 1981). In
all Timan areas, deeper and shallower shelf areas alternated and
facies varied from reefal to neritic cephalopod (House et al.,
2000). The Lyaiol’ community is regarded as a deeper-facies
analogue of the Vetlasyan-Sirachoy + Ukhta communities (Esin
et al., 2000) (Table 3).
The Voronezh, Evlanovo, and Livny communities of the CDF
lived in a shallow-water sea with normal salinity (Rodionova et
al., 1995) and are characterised by the absence of provincial
endemics (Table 3). No key taxa, such as psammosteids and
porolepiforms, are recorded from this district. Arthrodire and
ptyctodont placoderms dominated the communities, antiarchs
and acanthodians being a rare exception. Dipnoans are few
in number with a limited number of genera; Palaedaphus,
known otherwise from the Famennian of Belgium (Cloutier
and Candilier, 1995) is thus a local endemic for the Frasnian
in the CDF. Material originally identified as? Plourdos-
teus (Obrucheva and Obrucheva, 1977) from the Voronezh
Regional Stage seems to represent a new genus of the family
Plourdosteidae.
Another assemblage of Early rhenana Zone (Voronezh time)
is known from the Soligalich borehole (northeast of the Moscow
syneclise). With Psammosteus, Bothriolepis, and Holoptychius,
this grouping is close to that from the MDF, showing an open
faunal connection to the western part of Baltica. Bothriolepis
and Holoptychius were obtained from the Evlanovo deposits
in the Lyubim borehole of the neighbouring Yaroslavl’ Region
(Tikhomirov, 1967), also being in accordance with the vertebrate
assemblages of the MDF.
The provincial characteristics are based upon the only psam-
mosteid Obruchevia known only from the Pamu¯sˇis-Stipinai
Author's personal copy
11
2
O
.A
.L
eb
ed
ev
et
a
l.
/P
a
la
eo
wo
rld
19
(20
10
)1
08
–1
28
Table2
MiddleFrasnianvertebratecommunitiesoftheBalticaProvince.Basedupon:ObruchevaandObrucheva(1977),Luksˇevicˇs(1999),Esinetal.(2000),ValiukevicˇiusandKruchek(2000),Ivanovetal.(2006),
Moloshnikov(2008)andunpublishedauthor’sdata.
LocalendemicsProvincialendemicsQuasiendemics/didemicsPolydemicsCosmopolitansE-Cindex
DubnikcommunityofMDF(punctataZone)
Parapanderichthys“Acanthodes”
Jarvikina
Psammosteus
Plourdosteus
Asterolepis
Gyroplacosteus
“Rhynchodus”
“Dipterus”
“Ptyctodus”
Holonema
Eastmanosteus
Bothriolepis
Persacanthus
Holoptychius
Rhinodipterus
Moythomasia
1:2:3:3:8
DaugavacommunityofMDF(EarlyhassiZone)
Parapanderichthys“Acanthodes”
Jarvikina
Psammosteus
Plourdosteus
Eusthenopteron
“Rhynchodus”“Ptyctodus”
Holonema
Eastmanosteus
Bothriolepis
Holoptychius
1:2:3:1:5
SnezhacommunityofMDF(jamieaeZoneinterval)
PlatycephalichthysPsammosteus
Plourdosteus
Devononchus
Holonema
Bothriolepis
Holoptychius
1:0:3:0:3
SemilukicommunityofCDF(punctata-EarlyhassiZoneinterval)
GeptolepisPsammosteus
Plourdosteus
“Rhynchodus”“Ptyctodus”
Holonema
Holoptychius
Moythomasia
1:0:2:1:4
DomanikcommunityofSouthTiman(punctata-jamieaeZoneinterval)
Timanosteus
cf.Cheiracanthus
cf.Rhadinacanthus
“Acanthodes”Plourdosteus
Devononchus
“Rhynchodus”
Grossilepis
Eastmanosteus
Moythomasia
3:1:2:2:2
KraypolecommunityofMiddleTiman(punctata-jamieaeZoneinterval)
“Acanthodes”
Jarvikina
Psammosteus
Plourdosteus
Devononchus
Grossilepis“Ptyctodus”
Bothriolepis
Persacanthus
Holoptychius
Moythomasia
0:2:3:1:5
Ust’-BezmoshitsacommunityofNorthTiman(?)
JarvikinaPsammosteus
Asterolepis
GlyptolepisHolonema
Bothriolepis
Holoptychius
0:1:2:1:3
Middle-LateMatusevichcommunityofSevernayaZemlya(?)
Psammosteus
Asterolepis
Eusthenopteron
?Eastmanosteus
Bothriolepis
Holoptychius
0:0:3:0:3
Author's personal copy
O.A. Lebedev et al. / Palaeoworld 19 (2010) 108–128 113
Fig. 2. Distribution of vertebrate communities of the Baltica Province during the late Frasnian. Black squares indicate the most important vertebrate localities. Arrows
illustrate faunal interchanges between districts; abbreviations by the arrows list the main vertebrate groups participating in the dispersal. For abbreviations see Fig. 1.
community of the MDF and Kamenniy community of the North
Timan, showing a wide range of dispersal but known yet in
Baltica only from these areas (Table 3).
Two types of intra- and interprovincial connections are
registered for the late Frasnian of the Baltica (Fig. 2): (1) a con-
tinental, brackish- or even freshwater connection exemplified
by psammosteids (Psammosteus, Obruchevia), acanthodians
(Devononchus), and antiarchs (Bothriolepis); (2) presumably
shallow-water marine, possibly even hypersaline, realised by
the ptyctodonts (“Ptyctodus”, “Rhynchodus”), struniiforms
(Strunius), and dipnoans (Conchodus, Ganorhynchus). The sec-
ond type makes clear the presence of some didemics, polydemics
and cosmopolitans in the Central Russian faunas, their isola-
tion from the penetration of provincial endemics, typical for the
rest of Baltica. The presence of the elasmobranch Phoebodus,
known otherwise only from the continental shelves, supports
this assumption.
Three types of faunas may be established for the late Frasnian
within Baltica: (1) Fennoscandian (exemplified by Psammos-
teus, Obruchevia, and Devononchus), (2) marginal Baltican
(exemplified by presence of Conchodus and, in contrast, miss-
ing Holoptychius, which is widely spread in the other parts of
Baltica, but is also absent from the Armorican territories), and
(3) mixed faunas of Timan, which include the elements of both
previous faunas.
3.2. Famennian
3.2.1. Early Famennian (triangularis-rhomboidea time
interval) (Table 4, Fig. 3)
Early Famennian communities are documented from the
same districts as previously, but in Severnaya Zemlya the verte-
brate record is the last in the Devonian section, and no vertebrates
of this age are known from Middle and North Timan. A marine
Author's personal copy
11
4
O
.A
.L
eb
ed
ev
et
a
l.
/P
a
la
eo
wo
rld
19
(20
10
)1
08
–1
28
Table3
LateFrasnianvertebratecommunitiesoftheBalticaProvince.Basedupon:Esinetal.(2000),ValiukevicˇiusandKruchek(2000),Krupina(2004),MatukhinandMenner(1999),Moloshnikov(2002),Obruchev
(1959),Obrucheva(1962),Zakharenko(2007)andunpublishedauthor’sdata.
LocalendemicsProvincialendemicsQuasiendemics/didemicsPolydemicsCosmopolitansE-Cindex
Pamu¯sˇis-StipinaicommunityofMDF(rhenanaZone)
Walterilepis
Platycephalichthys
Obruchevichthys
ObrucheviaPsammosteus
Asterolepis
Devononchus
Grossilepis
“Dipterus”
“Ptyctodus”
Bothriolepis
Holoptychius
3:1:3:2:3
AmulacommunityofMDF(linguiformisZone)
Psammosteus
Devononchus
Bothriolepis
Holoptychius
0:0:2:0:2
VoronezhcommunityofCDF(EarlyrhenanaZone)
“Plourdosteus”
Ganorhynchus
“Rhynchodus”“Ptyctodus”
Bothriolepis
0:0:2:1:2
EvlanovocommunityofCDF(LaterhenanaZone)
Omalosteus
Palaedaphus
Conchodus“Ptyctodus”
Phoebodus
Strunius
2:0:1:0:3
LivnycommunityofCDF(linguiformisZone)
“Rhynchodus”“Ptyctodus”0:0:0:1:1
Vetlasyan-SirachoycommunityofSouthTiman(EarlyrhenanaZone)
“Acanthodes”
cf.Cheiracanthus
cf.Rhadinacanthus
Psammosteus
Devononchus
Conchodus
Holonema
Eastmanosteus
“Ptyctodus”
Bothriolepis
Holoptychius
Moythomasia
0:3:3:0:6
UkhtacommunityofSouthTiman(Laterhenana-linguiformisZoneinterval)
“Acanthodes”
cf.Cheiracanthus
cf.Rhadinacanthus
Psammosteus
“Dinichthys”
Devononchus
Glyptopomus
Eastmanosteus
Bothriolepis
Holoptychius
0:3:4:0:3
Lyaiol’communityofSouthTiman(Earlyrhenana-linguiformisZoneinterval,deep-wateranalogueoftheVetlasyan-SirachoyandUkhtacommunities)
“Acanthodes”Psammosteus
?Brachydeirus
Devononchus
Laccognathus
ProtacrodusEastmanosteus
Bothriolepis
Phoebodus
Moythomasia
0:1:4:1:4
KamenniyRucheycommunityofMiddleTiman(EarlyrhenanaZone)
PsammosteusGlyptolepisHoloptychius0:0:1:1:1
KamenniycommunityofNorthTiman(EarlyrhenanaZone)
ObrucheviaPlourdosteusHolonema
Bothriolepis
0:1:1:0:2
VavilovcommunityofSevernayaZemlya(?)
?PsammosteusBothriolepis
Holoptychius
0:0:1:0:2
Author's personal copy
O.A. Lebedev et al. / Palaeoworld 19 (2010) 108–128 115
Table 4
Early Famennian vertebrate communities of the Baltica Province. Based upon: Obruchev (1958), Vorobyeva (2004), Obrucheva, E.D. (1983), Obrucheva and
Obrucheva (1977), Ivanov and Luksˇevicˇs (1994), Matukhin and Menner (1999), Esin et al. (2000), Valiukevicˇius and Kruchek (2000), Moloshnikov (2001, 2004,
2008), Beznosov (2009), Beznosov et al. (2004) and unpublished author’s data.
Local endemics Provincial endemics Quasiendemics/didemics Polydemics Cosmopolitans E-C index
Eleja community of MDF (triangularis Zone)
Devononchus Holoptychius Bothriolepis
“Acanthodes”
0:1:0:1:2
Jonisˇkis community of MDF (crepida Zone)
Devononchus Holoptychius Phoebodus
Ctenacanthus?
Moythomasia?
0:1:0:1:3
Kursa community of MDF (rhomboidea Zone)
Haplacanthus
Cheiracanthus
Devononchus
?Glyptolepis Phyllolepis
Holoptychius
Bothriolepis
Protacrodus
0:3:1:2:2
Zadonsk community of CDF (crepida Zone)
Livnolepis
Rossolepis
Chelyophorus
Megapomus
Chirodipterus
Conchodus
Jakubsonia
Devononchus “Dinichthys”
Glyptopomus
Strunius
Holoptychius
Dipterus
Remigolepis
Bothriolepis
Protacrodus
7:1:3:2:3
Elets community of CDF (rhomboidea Zone)
Eunemacanthus Dipterus 1:0:0:1:0
Savinobor-Sosnogorsk community of South Timan (triangularis Zone)
cf. Holodipterus Haplacanthus
Devononchus
“Cheiracanthus”
Holoptychius Bothriolepis
“Acanthodes”
Moythomasia?
1:3:0:1:3
Malyutka community of Severnaya Zemlya (?)
Bothriolepis 0:0:0:0:1
transgression started in the early Famennian, moving from the
southeast to the northwest. In the MDF the sedimentation of tri-
angularis Zone (Eleja time) took place within a restricted shelf,
and even in the sabkha, gradually changing into a shallow-water
marine environment in the course of the transgression. Dur-
ing the later crepida Zone (Jonisˇkis time), carbonate and clay
muds were formed in a shallow sea with normal salinity, fol-
lowed by rhomboidea Zone (Kursa time), also characterised by
shallow-water marine conditions (Savvaitova, 1977) (Table 4;
Fig. 3).
Terrigenous material actively washed into the Moscovian Sea
from the Voronezh Peninsula, as well as from Fennoscandia
(Fig. 3), formed large subaquatic deltas during crepida Zone
(Zadonsk time), but eastwards the sedimentation type indi-
cates normal marine salinity. The connection of the CDF basin
to the Baltic syneclise was blocked by the Latvian Isthmus
(Rodionova et al., 1995), but in comparison to the late Frasnian
the provincial endemics Devononchus, polydemics Holopty-
chius and cosmopolitan Bothriolepis appeared in this part of
Baltica, suggesting a direct faunistic contact. The dominating
element in the Zadonsk community, Livnolepis zadonica (H.
Obrucheva, 1983) was also found in the Tver’ Region of Rus-
sia (Moloshnikov, 2004, 2008), extending the dispersal of this
fauna to the northwest. The intermediate location of this ver-
tebrate assemblage (Bothriolepis and Dipterus) on the way to
Timan was recorded in the Lyubim borehole (Yaroslavl’ Region:
Tikhomirov, 1967). During the successive rhomboidea Zone
(Elets time) the sediments were mostly formed in the normal
marine environments (Rodionova et al., 1995).
The Savinobor community of South Timan (triangularis
zone) lived in shallow shelf environments (House et al., 2000),
and the Sosnogorsk community (triangularis-crepida Zone
interval) occupied shallow-water lagoonal conditions that peri-
odically dried out (Beznosov et al., 2004; Beznosov, 2009).
As during the late Frasnian, the Latvian Isthmus played an
important role as a zoogeographical barrier, but became more
“transparent”, letting pass some vertebrates and obstructing tran-
sit for others. This is exemplified by the provincial endemic
Devononchus, known from all major Baltica districts, and the
biogeographically significant placoderm Phyllolepis, which did
not penetrate to the CDF from the MDF. On the contrary, the
Famennian cosmopolitan Remigolepis, appearing in Baltica for
the first time in the CDF, remains unknown in the MDF. How-
ever, both genera coexist and were closely associated in East
Greenland (Blom et al., 2007). A direct Baltica–Greenland
connection would require either recognition of the freshwa-
ter environments as habitats for these vertebrates, and their
capability of fluvial dispersal, or recognition of a western pas-
sage to Greenland through Scotland. At the same time, the
acanthodian type of connection within Baltica is definitely non-
Author's personal copy
116 O.A. Lebedev et al. / Palaeoworld 19 (2010) 108–128
Fig. 3. Distribution of vertebrate communities of the Baltica Province during the early Famennian. Black squares indicate the most important vertebrate localities.
Arrows illustrate faunal interchanges between districts; abbreviations by the arrows list the main vertebrate groups participating in the dispersal. For abbreviations
see Fig. 1.
marine during the Famennian (in contrast to the late Frasnian
and middle–late Eifelian; Luksˇevicˇs et al., 2010). It remains
unclear, however, whether the MDF–Timan connection charac-
terised by acanthodians, Holoptychius and Bothriolepis, was a
direct one, or used coastal pathways by the southern coast of
Baltica.
The low-diversity communities of the MDF and South Timan
are characterised by complete or almost complete absence of
local endemics (Table 4). In contrast, in the Zadonsk commu-
nity of the CDF local endemics dominate. The characteristic
provincial endemics include the acanthodians Haplacanthus,
Cheiracanthus, and Devononchus, comparable to the situation
in the early Frasnian.
There are no records of the dipnoans Chirodipterus and Con-
chodus elsewhere in the Famennian; most probably the CDF was
a refugium for these fishes. The few didemics and polydemics
demonstrate predominantly Laurentian affinities.
3.2.2. Middle Famennian (marginifera-trachytera time
interval) (Table 5)
The middle Famennian communities continued their exis-
tence in the MDF, CDF, and South and North Timan. During
marginifera Zone (Akmene time) in the MDF, only a few verte-
brates are known from almost normal marine environments. In
the following trachytera Zone (Spa¯rnene time), the shallow sea
gradually regressed, with the formation of shallow-water coastal
deposits, including a richer vertebrate assemblage as well as
plant remains (Savvaitova, 1977). In the CDF the marginifera
Zone (Lebedyan’ time) was characterised by normal marine and
hypersaline lagoonal aquatic environments. Limited water input
continued from the east and south-east; the connection to the
Baltic syneclise was still blocked by the Latvian Isthmus; dur-
ing the trachytera Zone (Optukha time), the territory of the CDF
was occupied by a shallow-water lagoonal basins characterised
by increased salinity (Rodionova et al., 1995) (Table 5).
Author's personal copy
O.A. Lebedev et al. / Palaeoworld 19 (2010) 108–128 117
Table 5
Middle Famennian vertebrate communities of the Baltica Province. Based upon: Obrucheva, O.P. (1983), Obrucheva and Obrucheva (1977), Esin et al. (2000),
Valiukevicˇius and Kruchek (2000), Luksˇevicˇs (2001) and unpublished author’s data.
Local endemics Provincial endemics Quasiendemics/didemics Polydemics Cosmopolitans E-C index
Akmene community of MDF (marginifera Zone)
Megapomus Chelyophorus ?Glyptolepis Phyllolepis
Holoptychius
Bothriolepis 1:1:1:2:1
Spa¯rnene community of MDF (∼trachytera Zone)
Platycephalichthys
Cryptolepis
Conchodus
Chelyophorus
Devononchus
Homacanthus
?Glyptolepis
Phyllolepis
“Dunkleosteus”
Holoptychius
Bothriolepis
Protacrodus
“Acanthodes”
3:2:2:3:3
Lebedyan’ community of CDF (marginifera Zone)
Chelyophorus Holoptychius
Dipterus
0:1:0:2:0
Optukha community of CDF (trachytera Zone)
Dipterus 0:0:0:1:0
Ust’-Pechora community of South Timan (marginifera-trachytera Zone interval)
cf. Cheiracanthus
cf. Rhadinacanthus
Devononchus “Acanthodes” 2:1:0:0:1
Pokayama community of North Timan (marginifera Zone)
Phyllolepis
Holoptychius
Bothriolepis 0:0:0:2:1
The provincial endemics Chelyophorus and Devononchus
characterise the middle Famennian communities, although both
are absent from North Timan, and the latter is unknown from
the CDF (Table 5). The CDF assemblage is impoverished pos-
sibly due to unfavourable salinity in the basin; in South Timan
only poorly identified acanthodian scales are recorded, so no
zoogeographical conclusions are possible. The presence of the
placoderm Phyllolepis in the Baltic syneclise and North Timan
marks the highest point of its dispersal in Laurussia.
3.2.3. Late Famennian (postera-expansa time interval)
(Table 6, Fig. 3)
Carbonate deposits were formed in a shallow-water sea with
normal salinity during postera Zone (Piemare time) in the MDF,
with sandy deposits closer to the shoreline. During the following
expansa Zone (Ketleri time), deposition took place in a low-tidal
near-shore environment (Luksˇevicˇs and Zupin¸sˇ, 2004). During
postera-expansa Zone interval (Plavsk time), mostly lagoonal
basins occupied the CDF territory, interrupted by locally formed
Table 6
Late Famennian vertebrate communities of the Baltica Province. Based upon: Obruchev (1958), Obrucheva, O.P. (1956, 1962, 1983), Obrucheva and Obrucheva
(1977), Krupina (1986, 1999, 2000, 2004), Lebedev (1983, 1995), Lebedev and Luksˇevicˇs (1996), Esin et al. (2000), Valiukevicˇius and Kruchek (2000), Vorobyeva
(2004), Luksˇevicˇs and Zupin¸sˇ (2004), Moloshnikov (2008) and unpublished author’s data.
Local endemics Provincial endemics Quasiendemics/didemics Polydemics Cosmopolitans E-C index
Piemare community of MDF (postera Zone)
Platycephalichthys Chelyophorus “Dinichthys”
Homacanthus
?Glyptolepis
Bothriolepis 1:1:3:0:1
Ketleri community of MDF (expansa Zone)
Ventastega Devononchus
“Cheiracanthus”
Cryptolepis
Ventalepis
Orlovichthys
Glyptopomus Holoptychius Bothriolepis
“Acanthodes”
1:5:1:1:2
Plavsk community of CDF (postera-Early expansa Zone interval)
Tuberospina
Pycnacanthus
Onychodus
Holodipterus
Grossipterus
Chirodipterus
Conchodus
Chelyophorus
Devononchus
“Cheiracanthus”
Cryptolepis
Ventalepis
Orlovichthys
“Dinichthys”
Glyptopomus
Glyptolepis
Strunius
Jarvikia
Holoptychius
Dipterus
Bothriolepis
“Acanthodes”
Moythomasia
7:6:5:2:3
Dzhebol’ community of South Timan (postera Zone)
Devononchus “Acanthodes” 0:1:0:0:1
Author's personal copy
118 O.A. Lebedev et al. / Palaeoworld 19 (2010) 108–128
Fig. 4. Distribution of vertebrate communities of the Baltica Province during the late Famennian. Black squares indicate the most important vertebrate localities.
Arrows illustrate faunal interchanges between districts; abbreviations by the arrows list the main vertebrate groups participating in the dispersal. For abbreviations
see Fig. 1.
river estuaries. These two types of sedimentary environments are
bound to each other by a gradual facies transition (Rodionova et
al., 1995) (Table 6).
Correlation of the Plavsk Regional Stage and especially its
Oryol-Saburovo beds to the upper part of the Ketleri Regional
Stage in Latvia (MDF) is controversial as shown by Lebedev and
Luksˇevicˇs (1996); the accepted dating by conodonts indicates
the latter is younger than the former. However, the vertebrate
assemblages differ only insignificantly (numerous dipnoans and
struniiforms in the former versus the tetrapod Ventastega in the
latter). This may be explained by the wider temporal range of a
single long-lived vertebrate fauna in both areas of the Baltica
Province, most probably within the continental basins. This
problem is aggravated by the absence of a clear aquatic connec-
tion of the basin in western Latvia to the CDF (Fig. 4). Isolated
Dipterus finds showing the extension of the CDF basin were
mentioned by Tikhomirov (1967) from deposits of the same age
in the Kostroma and Yaroslavl’ Regions. “Dinichthys” mach-
laevi Obrucheva (1956) from the CDF is based upon poorly
diagnostic material and needs restudy; “Cheiracanthus” scales
from the Plavsk and Ketleri communities await description, and
possibly belong to a new genus.
The provincial endemic assemblage includes the character-
istic ptyctodont Chelyophorus, acanthodians Devononchus and
“Cheiracanthus”, the osteolepiform Cryptolepis, the porolepi-
form Ventalepis, and the dipnoan Orlovichthys. Chelyophorus
persists from the middle Famennian, and the acanthodian
Devononchus characterises Baltica since the middle Givetian.
The remaining sarcopterygians distinguish the Ketleri-Plavsk
fauna (Table 6).
The Plavsk vertebrates show the dominance of local and
provincial endemics while in the Ketleri community provincial
endemics definitely prevail. This may indicate stronger isola-
tion of the CDF fauna in combination with more favourable
Author's personal copy
O.A. Lebedev et al. / Palaeoworld 19 (2010) 108–128 119
Table 7
The Latest Famennian vertebrate communities the Baltica Province. Based upon: Savvaitova (1977), Lebedev (1992), Krupina (1987), Luksˇevicˇs (1991), Alekseev
et al. (1994).
Local endemics Provincial endemics Quasiendemics/didemics Polydemics Cosmopolitans E-C index
ˇSk¸ervelis community of MDF (∼Late expansa Zone)
Holoptychius 0:0:0:1:0
Khovanshchina community of CDF (Late expansa Zone)
“Cheiracanthus”
Devononchus
“Chrysolepis”
Tulerpeton
Strunius Holoptychius
Eusthenodon
Andreyevichthys
Remigolepis
Bothriolepis
“Acanthodes”
Moythomasia
4:0:1:3:4
environmental conditions. Didemics and polydemics emphasise
Laurentian affinities (see Sections 4.2.1 and 4.2.2; Fig. 4). Mid-
dle Famennian endemism in the CDF is not clearly expressed,
whereas the early and late Famennian show strong prevalence
of local and provincial endemics.
3.2.4. Latest Famennian (Late expansa time interval)
(Table 7)
By the end of the Famennian the habitat for aquatic ver-
tebrates within the Baltica Province was strongly reduced to
small areas in the MDF and CDF. During the Late expansa
Zone ( ˇSk¸ervelis time) the restricted area in the MDF was a
shallow-water partly isolated basin (Savvaitova, 1977), but later
its sediments were reworked under subaerial conditions of an
arid climate (Stinkulis, 2008) (Table 7).
No vertebrates are recorded from the Ozerki deposits (Mid-
dle expansa Zone), possibly because of the hypermineralized
nature of the basins during this time in the central part of the Rus-
sian Platform. During Khovanshchina time (Late expansa Zone),
sedimentation occurred in near-shore environments under con-
ditions of variable salinity. This basin was influenced by the
sea in the south-east of the platform, and by freshwater input
from the northern slope of the Voronezh Island (Rodionova et
al., 1995; Alekseev et al., 1994).
No provincial endemics may be suggested for this age
because it is impossible to compare with other districts (Table 7).
Most taxa fall within the polydemic and cosmopolitan cate-
gories, showing global, especially Gondwanan, affinities (see
Sections 4.2.2 and 4.2.3) and good environmental conditions
for faunal interchange.
Two species of a new osteolepidid genus, designed here
as “Chrysolepis” (sensu Lebedev, 1992), are referred to local
endemics.
4. Distribution of didemics/quasiendemics, polydemics,
and cosmopolitans within the Baltica Province and
globally
This section considers the vertebrate genera found within
the Baltica Province classified into the categories of
didemics/quasiendemics, polydemics, and cosmopolitans by
comparison to their distribution in the other areas of the world
during the Late Devonian.
4.1. Frasnian
4.1.1. Didemics/quasiendemics
The psammosteid Psammosteus, apart from the Russian Plat-
form, is known from Scotland, Canadian Arctic Archipelago,
and Severnaya Zemlya (Blieck et al., 2002), and Psammolepis
from Arctic Canada (Elliott et al., 2003).
The arthrodire Plourdosteus is recorded from the west-
ern margin of the Russian Platform (Main Devonian Field:
Obrucheva, 1962; and Holy Cross Mountains, Poland:
Kulczycki, 1957) and Canada (Denison, 1978). A specimen from
Bergisch Gladbach, Germany is attributed to this genus with
reservations (Denison, 1978), so further study might require the
attribution of this genus to the quasiendemics/didemics.
The widespread antiarch Asterolepis occurs in New York
State and Nevada (USA), Scotland, and Belgium (Denison,
1978). Its identification in possible Frasnian of Columbia
(Janvier and Villarroel, 2000) would indicate connections
between Euramerica and Western Gondwana.
Apart from Baltica, the porolepiform Laccognathus is known
from Canada (Daeschler et al., 2003) and the osteolepiform
Eusthenopteron from Canada and Greenland (Schultze and
Cloutier, 1996).
The Frasnian dipnoan Ganorhynchus is recorded from
North America (Newberry, 1889) and Belgium (Cloutier and
Candilier, 1995);Conchodus is known from Poland and Scotland
(Kulczycki, 1957; Vorobyeva and Obruchev, 1964).
In comparison to Givetian, the diversity of the didemics
remains almost the same. These genera show almost exclusively
east Laurentian connections, as during the Middle Devonian
(Table 8).
4.1.2. Polydemics
The dental plates of “Rhynchodus” are known from the
Holy Cross Mountains (Poland), New York State (USA),
and Bad Wildungen (Germany). Apart from the MDF, the
arthrodire Gyroplacosteus was described from the Kuznetsk
Basin (Siberia), Poland, and Germany (Denison, 1978).
The antiarch Grossilepis is rather widely distributed in the
MDF; another species of this genus has been reported from Scot-
land (Miles, 1968) and Siberia (Obruchev and Sergienko, 1960),
but the latter was based on limited material and needs restudy.
Apart from Baltica, the porolepiform Glyptolepis is known
from Scotland (Ahlberg, 1989), Canada (Schultze and Cloutier,
Author's personal copy
120 O.A. Lebedev et al. / Palaeoworld 19 (2010) 108–128
Table 8
Zoogeographic evolution of non-endemic Late Devonian vertebrates and their faunistic interchanges within the Baltica faunistic Province.
Time Zoogeographic categories Vertebrate groups Prevailing interconnections
Famennian Quasiendemics/didemics Acanthodians
porolepiforms
struniiforms
dipnoans
Laurentia
Polydemics Phyllolepids
porolepiforms
osteolepiforms
dipnoans
Laurentia
Siberia
East
Gondwana
Cosmopolitans Antiarchs
chondrichthyans
actinopterygians
Laurentia
Siberia
East Gondwana
North China
South China
Frasnian Quasiendemics/didemics Psammosteiforms
arthrodires
antiarchs
elasmobranchs
porolepiforms
osteolepiforms
dipnoans
Laurentia
Polydemics Ptyctodonts
arthrodires
antiarchs
acanthodians
porolepiforms
osteolepiforms
struniiforms
dipnoans
Laurentia
Siberia
Cosmopolitans Ptyctodonts
arthrodires
antiarchs
struniiforms
actinopterygians
Laurentia
North Gondwana
East Gondwana
Siberia
Armorica
1996), and Minusa (Obruchev, 1941). The last find is based upon
a single scale and needs revision.
The diversity of polydemics increased significantly after
the Givetian; antiarchs, acanthodians, and various sarcoptery-
gians were added to the list (Table 8; see also Luksˇevicˇs et
al., 2010). The distribution of the majority of the polydemic
genera included the eastern part of Laurentia and Armorica.
Gyroplacosteus, Grossilepis, and Glyptolepis show connections
to Siberia. The increase of the polydemics may reflect the general
Frasnian increase of diversity (Long, 1993).
4.1.3. Cosmopolitans
The Middle-Upper Devonian ptyctodont placoderm “Ptycto-
dus” is very widely distributed and especially abundant in the
shallow-water marine deposits during the Frasnian. It is known
from numerous localities in Baltica, Laurentia, East and North
Gondwana, and Siberia (Denison, 1978).
The distribution of the arthrodire Holonema during the
Frasnian includes Laurentia: Eastern USA, Canadian Arc-
tic Archipelago, Scotland; North Gondwana: Iran, Morocco,
Afghanistan, Turkey, and East Gondwana: Australia (Schultze,
1973; Denison, 1978; Lelièvre et al., 1993; Janvier et al.,
2007).
The antiarch Bothriolepis is almost globally distributed
in Laurentia: Scotland, Canada, and USA (Denison, 1978);
Siberia and adjacent regions (Tuva, Minusa, Kuznetsk Basin,
Tien-Shan: Obruchev, 1955; Denison, 1978), Kotel’ny Island
(Mark-Kurik, 1974); Kazakhstan (Malinovskaya, 1992); South
and North China (Zhu, 2000), and Gondwana: Australia and
Antarctica (Young and Long, 2005), Venezuela (Young et
al., 2000) and Turkey (Janvier, 1983). Some cosmopolitan
species of this genus demonstrate connections between vari-
ous areas within the Old Red Sandstone continent even on the
specific level; for example, Bothriolepis cellulosa and B. pan-
deri closely resemble B. canadensis from Canada (Luksˇevicˇs,
2001).
The phoebodontiform elasmobranch Phoebodus is very
widely distributed during the Frasnian, but its only epicra-
tonic species Ph. bifurcatus apart from the CDF is known
from Bashkortostan, Timan, and Poland in the Baltica shelves,
and in the Armorican Province (Moravia) (Ginter and Ivanov,
1992).
Various species of the acanthodian Atopacanthus were dis-
covered in USA and Germany (Denison, 1979); those of the
closely related genus Persacanthus occur in Iran, Nevada, and
Timan (Beznosov, 2002).
Author's personal copy
O.A. Lebedev et al. / Palaeoworld 19 (2010) 108–128 121
Holoptychius was described from Laurentia (Scotland:
Ahlberg, 1989, 1992; Canada: Schultze and Cloutier, 1996 and
USA: Ørvig, 1957); North Gondwana (Iran: Schultze, 1973),
and West Gondwana (Columbia), but this last identification is
dubious (Janvier and Villarroel, 2000).
The osteolepiform Latvius has been reported from Germany
(Jessen, 1966) and New Brunswick, Canada (Greiner, 1977);
the struniiform Strunius from Poland (Ginter, 2002), Germany
(Jessen, 1966), and Iran (Janvier and Martin, 1979). The dipnoan
Rhinodipterus was discovered in Germany, France, and Belgium
(Jessen, 1966; Cloutier and Candilier, 1995), and Griphognathus
in Germany (Gross, 1956), Nevada (Schultze and Cloutier, 1996)
and Australia (Miles, 1977; Andrews et al., 2006).
During the Frasnian the struniiform Onychodus was widely
spread in North and East Gondwana (Iran: Janvier and Martin,
1979; Afghanistan: Blieck et al., 1982; Turkey: Lelièvre et al.,
1993; Australia: Andrews et al., 2006), Laurentia (USA: Elliott
et al., 2000; northern France: Cloutier and Candilier, 1995) and
Armorica (Germany: Jessen, 1966).
The actinopterygian Moythomasia is spread in Armor-
ica (Germany: Gross, 1950) and East Gondwana (Australia:
Gardiner, 1984).
The cosmopolitans, including ptyctodonts, arthrodires
(Holonema), antiarchs, struniiforms, and actinopterygians,
demonstrate the maximum diversity in this category during the
Middle-Late Devonian (Table 8). Holonema changed its sta-
tus from polydemic to the cosmopolitan; the last three groups
remain cosmopolitan since the Givetian. Most of the cosmopoli-
tan connections indicate to Laurentia, Armorica, East and North
Gondwana.
4.2. Famennian
4.2.1. Didemics/quasiendemics
Apart from the Baltica Province, the acanthodian Homacan-
thus was recorded from Laurentia (Ohio, USA: Denison, 1979).
The Famennian record of the porolepiform Glyptolepis
includes localities in Laurentia (France: Cloutier and Candilier,
1995). Apart from the Baltica, Glyptopomus is known from
Scotland and Belgium (Jarvik, 1950; Cloutier and Candilier,
1995); Strunius from Colorado, USA (Ginter, 2001). The dip-
noan Jarvikia is also known from East Greenland (Blom et al.,
2007).
The proportion of didemics strongly decreased in compari-
son to the Frasnian (Table 8). Arthrodires, chondrichthyans, and
antiarchs shifted to the polydemic and cosmopolitan categories.
The remaining genera show Laurentian affinities.
4.2.2. Polydemics
The distribution of the phyllolepid placoderm Phyllolepis
and its relationships to the Famennian zoogeography was earlier
mentioned by Lebedev (1985, 2004) and extensively discussed
by Young (2006). It was demonstrated that this placoderm is
closely associated to the earliest tetrapods. During the Famen-
nian, this fish spread to Laurussia (East Greenland, Scotland,
Belgium, Pennsylvania, USA, Latvia; North Timan) from Gond-
wana (Young, 2006).
Beyond the limits of the Baltica Province the porolepiform
Holoptychius is known from Laurentia (Greenland, Scotland,
Belgium, France: Jarvik, 1972; Cloutier and Candilier, 1995),
East Gondwana (Australia: Johanson and Ritchie, 2000) and
Siberia (Kuznetsk Basin: Ivanov and Rodina, 2004).
The tristichopterid Eusthenodon was previously described
from Laurentia: Greenland (Blom et al., 2007), Pennsylvania
(Elliott et al., 2000), Belgium (Clément, 2002), and East Gond-
wana (Australia: Johanson and Ritchie, 2000). It may be present
in the Famennian of South Africa, but still awaits description
(Anderson et al., 1999).
Numerous finds attributed to the genus Dipterus from North
America (Eastman, 1907) are dubious and require revision
(Friedman and Daeschler, 2006); for the time being the sta-
tus of this material remains unchanged. Dipterus finds from
France and Belgium were summarised by Cloutier and Candilier
(1995). Obruchev (1940,1960) presented materials from Siberia
(Kuznetsk Basin).
Anderson et al. (1994, 1999) illustrated an isolated paras-
phenoid very similar to that of Andreyevichthys; despite
insufficiency of the material, this affinity seems obvious. Ginter
(2001) figured Andreyevichthys juvenile tooth plates from Col-
orado closely resembling those described by Krupina and Reisz
(1999). This genus is conditionally regarded here as polydemic;
further studies in South Africa and westerm United States may
support or reject this opinion. In the latter case the genus would
become a didemic, or a local endemic.
Close connections of the Baltica Province to Laurentia
persisted during the Famennian (Table 8). The polydemics
demonstrate interrelationships with Siberia, as during the Fras-
nian, and East Gondwana. The South African link still needs to
be supported by further evidence.
4.2.3. Cosmopolitans
The antiarch Bothriolepis is one of the frequent cosmopolitan
elements in Famennian faunas. It is known from Laurentia: Scot-
land; Greenland; Pennsylvania, USA, Belgium (Denison, 1978);
Siberia: Kuzbass (Obruchev, 1960); East Gondwana (Australia:
Young, 1988); China (Zhu, 2000) and South Africa (Anderson
et al., 1994).
The antiarch Remigolepis is also widely distributed, being
found in Laurentia: East Greenland (Blom et al., 2007), Penn-
sylvania, USA (Daeschler et al., 2003); Scotland (Andrews,
1978); Baltica: Tula Region, Russia (Luksˇevicˇs, 1991); East
Gondwana: Australia (Johanson, 1997); North and South China:
Ningxia, Inner Mongolia, Hunan (Zhu, 2000) and ? Tarim:
Kyrghyzstan (Panteleyev, 1992). The last record requires age
refinement, being currently defined only as Upper Devonian.
Famennian records of the chondrichthyan Protacrodus are
numerous and globally distributed. Ginter (2001) regarded this
fish as an indicator of shallow-water marine biofacies. Similarly
dispersed genera Protacrodus and Phoebodus were spread in
the coastal shelf zones and epicratonic seas of Laurentia: Iowa
and Colorado, USA (Gross, 1973; Ginter, 2001), Canada (Ginter
and Turner, 1999); Baltica: Poland, Cisurals (Ginter and Ivanov,
2000), Belarus (Esin et al., 2000), Latvia (Ivanov and Luksˇevicˇs,
1994); former Armorican territories: Germany (Ginter, 1999),
Author's personal copy
122 O.A. Lebedev et al. / Palaeoworld 19 (2010) 108–128
France (Ginter, 2000) and Italy (Randon et al., 2007); East and
North Gondwana: Australia (Young and Turner, 2000), Morocco
(Derycke, 1992), Iran (Ginter et al., 2002); Siberia: Kuznetsk
Basin (Ivanov et al., 1992) and China: Guizhou and Gansu (Zhu,
2000). Various skeletal fragments of Ctenacanthus were found
in USA (Zangerl, 1981), Bolivia (Janvier, 2003) and Gansu,
China (Zhu, 2000).
Apart from Baltica the actinopterygian Moythomasia is
recorded in Laurentia (France: Rieman et al., 2002) and North
Gondwana (Afghanistan: Blieck et al., 1982; Iran: Long and
Hairapetian, 2000).
Only the antiarch Bothriolepis persisted as a cosmopolitan
after the Givetian. Remigolepis became a cosmopolitan as well,
but with a smaller area of distribution. The cosmopolitan cat-
egory became dominated by diversifying chondrichthyans, and
to a lesser extent actinopterygians (Table 8).
4.3. Pelagic and continental margin assemblages of the
Baltica Province periphery
Research of past decades has documented a number of the
Givetian-Famennian vertebrate assemblages in the pelagic facies
around the Baltica Province: North and South Urals and Poland
(e.g., Ivanov, 1999; Ginter and Ivanov, 2000; Ginter, 2001;
Ginter et al., 2002). These assemblages are composed mostly
of various chondrichthyan taxa belonging to Phoebodontidae,
Protacrodontidae, Omalodontidae, Stethacanthidae, and Sym-
moriidae. Isolated skeletal parts of acanthodians, struniiforms,
and actinopterygians are often found together. As shown by
Ginter (2000) and Ginter et al. (2002), these shark-dominated
assemblages occupied the continental margins of Laurentia,
Armorica, Baltica, and other provinces, and were largely absent
from continental or even near-shore communities, thus showing
no geographical subdivision. These authors proposed vertical
zoogeographical subdivision to distinguish between shallow-
and deep-water communities.
However, no separating line can be drawn between the con-
tinental and pelagic faunas, as members of one fauna could
penetrate to the habitat of another. This is exemplified by inva-
sion of some species of the chondrichthyans Phoebodus and
Protacrodus into shallow-water epicratonic basins of Baltica
during the late Frasnian of the CDF and Timan (Table 3), and
early Famennian of MDF and CDF (Table 4), during the periods
of high transgression.
Assigning such composite peripheral vertebrate commu-
nities to one or another province presents difficulties to a
palaeogeographer, for example the vertebrate faunas of the
Holy Cross Mountains (Poland). The absence of characteris-
tic provincial endemics precludes attribution of these faunas to
the Baltica Province. However, during the middle-late Givetian
the Baltican-Armorican didemic Conchodus and the Laurus-
sian didemic Glyptolepis entered the Holy Cross Mountains
communities (Kulczycki, 1957; Liszkowski and Racki, 1992;
Ivanov and Ginter, 1997). Again, during the middle Fras-
nian, the Laurussian didemics Psammosteus, Plourdosteus, and
Eusthenopteron are known. This may be explained by regres-
sions resulting in shallower seas, making possible the dispersal
of continental vertebrates from which they were earlier excluded
by deeper water. For the moment, the question on the affinity of
this faunal succession to a certain province remains unresolved.
5. General palaeozoogeographical tendencies during
the Late Devonian within the Baltica Province
5.1. Zoogeographical evolution of vertebrates
As shown in Table 8, the distribution of various verte-
brate groups by the zoogeographical E-C categories during
Late Devonian is uneven, perhaps due to ecological con-
strains of environments (e.g., salinity) or the nature of the
animals themselves (e.g., low swimming capabilities). The
agnathans (thelodonts and psammosteids), phyllolepid pla-
coderms, porolepiforms, osteolepiforms, and dipnoans never
enter the cosmopolitan category (see also Luksˇevicˇs et al.,
2010).
Other groups originally fall into the lowest E-C category,
then flourish (joining all categories), and decrease, like the acan-
thodians, arthrodires, antiarchs, ptyctodonts, and struniiforms.
The decline may be expressed by lowering of the category
status, as demonstrated by the last group. Others, like elasmo-
branchs and actinopterygians, attain their cosmopolitan maxima
by the Givetian-Famennian and pass into the Carboniferous. The
ingress of the phyllolepids during the Famennian already at the
high polydemic level supports Young’s (2003, 2006) idea on
their spreading to Laurussia from Gondwana.
All categories (quasiendemics/didemics, polydemics, and
cosmopolitans) attain their maxima during the Frasnian, cor-
responding to the increase of diversity demonstrated by Long
(1993). The quasiendemics/didemics demonstrate their minima
during the Famennian, the polydemics during the Givetian, and
the cosmopolitans during the Eifelian (Luksˇevicˇs et al., 2010).
The data on minima are not completely reliable, as each future
find would affect the presented result.
5.2. Faunal interchanges of the Baltica zoogeographical
Province in space and time
Faunal interchanges between the zoogeographical provinces
possibly depend on the proximity of the palaeocontinents, pres-
ence of physico-geographical barriers, and eustatic sea-level
changes (see Section 5.3). These interchanges may be inferred
from the interruptions of earlier established connections. As
shown in Table 8, the constant interrelationship between Baltica
and Laurentia is not surprising, given the placement of both
provinces over the same landmass, the Old Red Sandstone con-
tinent. The Armorican connection is also very stable until the
dissipation of this province in the Famennian (Lebedev and
Zakharenko, 2010), although the connections in the quasien-
demic/didemic category are missing.
Interchanges with East Gondwana appear in the Givetian and
stay at high (polydemic-cosmopolitan) levels into the Famen-
nian. The North Gondwanan connection appears only during
the Frasnian; the West Gondwanan one is almost absent.
Author's personal copy
O.A. Lebedev et al. / Palaeoworld 19 (2010) 108–128 123
Table 9
Distribution of the maxima of the E-C index in the vertebrate communities of the Baltica zoogeographical Province. Large circles—significant meanings,
small circles mark insignificant maxima or single records of taxa in the category. Empty boxes stand for missing deposits; labeled empty boxes denote
absence of vertebrates in the corresponding formations.
LE—local endemics, PE—provincial endemics, DI—didemics/quasiendemics; PO—polydemics, CO—cosmopolitans.
Author's personal copy
124 O.A. Lebedev et al. / Palaeoworld 19 (2010) 108–128
The connection to Siberia is interrupted during the Middle
Devonian, and restored in the Frasnian, but shows no close
quasiendemic/didemic associations. A distant South China link
is observed only during the Middle Devonian and Famennian in
the cosmopolitan category.
5.3. Communities structure, dispersal conditions and
sea-level changes
The E-C index based on the suggested endemic-cosmopolitan
distributional types presents various combinations. Its structure
reflects the isolation rate of the community as demonstrated by
the balance of local and provincial endemics, polydemics, and
cosmopolitans. Predominance of local and provincial endemics
suggests complete or relative isolation of the community not
only from the rest of the province, but also from the open sea.
The E-C index variation in time reflects changes in faunal iso-
lation/connections of the communities.
Alekseev et al. (1996) published a relative Devonian sea-level
curve for the Moscow syneclise, a part of the territory of the East
European platform. The analysis of the most important indices
(large dots, Table 9) demonstrates approximate correlation of
the E-C fluctuations with the sea-level curve.
During the Eifelian the depth of the basin increased. This
is reflected by predominance of polydemics and cosmopolitans
in the Pärnu, Narva, Mosolovo-Cherniy Yar and Lekkey-Yaga
communities.
The Givetian demonstrates a new, very unclearly expressed
cycle, and almost all communities show significant endemism-
didemism.
During the Frasnian a very strong transgression is interrupted
during jamieae time. The Pl¸avin¸as, Dubnik, and Daugava com-
munities of the MDF, as well as the Timan and Ust’-Yarega
communities of the Timan, correspondingly demonstrate strong
polydemism-cosmopolitanism (Tables 1 and 2). Unfortunately,
vertebrates of the late Frasnian are poorly known (Table 3),
and do not demonstrate a clear picture, showing concentration
of both didemics and cosmopolitans (except the Vetlasyan-
Sirachoy in the South Timan), although the same tendency is
suggested by the data.
In the Famennian the maximum transgression occurred in the
Zadonskian-Eletsian interval (crepida-rhomboidea zone inter-
val). In contrast, the Kursa and Zadonsk communities exemplify
well-pronounced endemism, possibly explained by isolation of
the former basin (Fig. 3) and special deltaic environments of
the latter. Subsequently the sea-level decreased (Fig. 4); cor-
respondingly the Ketleri-Plavsk fauna shows strong endemism
(Table 6). A minor pulse occurred during the Khovanshchinian,
as illustrated by cosmopolitanism and endemism at a time of
the corresponding community in the CDF (Table 7). However,
in this case, local endemism is possibly due to the special habi-
tat in the peculiar coastal environments of the epicontinental
sea.
Thus, certain dependence is proposed between the E-C com-
position of Devonian vertebrate communities and sea-level
changes. This observation correlates with the conclusions made
by Turner and Young (1997) for the East Gondwanan and Zhao
and Zhu (2007) for the Chinese vertebrate faunas. Deepening
would result in increased cosmopolitanism, whilst expansion of
endemism may be due to regressions, which increased isolation
of faunas, and perhaps created advantages for radiation within
limited areas.
6. Conclusions
Main results of the present study are summarised as follows:
• subdivisions of community members into E-C categories is
an effective tool in zoogeography;
• assemblages of provincial endemics permit assignment of
local communities to a particular zoogeographical province;
• dominance of local endemics suggests isolation of the com-
munity from the rest of the province, and also from the open
sea;
• two types of dispersal are proposed for Devonian aquatic ver-
tebrates: 1. normal marine and 2. continental, presumably
freshwater. The first is exemplified by acanthodians during
the Middle Devonian, and by ptyctodonts, struniiforms, and
some dipnoans during the Late Devonian; the second by
psammosteids, antiarchs, and porolepiforms during the Mid-
dle Devonian and by psammosteids, acanthodians, and some
arthrodires during the Late Devonian;
• E-C index variation in time reflects tendencies of faunal iso-
lation/connections of communities;
• constant faunal interchanges in the didemic/quasiendemic,
polydemic, and cosmopolitan categories are observed
between Baltica and Laurentia, but connections to other
provinces are sporadic;
• various vertebrate groups show different dispersal capa-
bilities. Agnathans (both thelodonts and psammosteids),
phyllolepid placoderms, porolepiforms, osteolepiforms, and
dipnoans never enter the cosmopolitan category;
• no clear line separates continental from pelagic faunas, as
the members of one fauna often penetrate the living area of
another;
• the E-C composition of Devonian vertebrate communities
generally correlates with sea-level changes. Transgressions
resulted in increased cosmopolitanism; augmentation of
endemism may be due to regressions causing greater isola-
tion. Separation from the main sea basin may have created
reduction of competition, causing radiation within limited
areas.
Acknowledgements
The authors are grateful for many of their colleagues, includ-
ing P. Ahlberg, A. Blieck, G. Clément, D. Goujet, W.J. Zhao
and M. Zhu for fruitful discussions of the problems of the
Devonian palaeozoogeography and for encouraging further
research in this field. G. Young and M. Zhu as reviewers made
numerous useful comments, which greatly improved the manu-
script.
Author's personal copy
O.A. Lebedev et al. / Palaeoworld 19 (2010) 108–128 125
References
Ahlberg, P.E., 1989. Paired fin skeletons and relationships of the fossil group
Porolepiformes (Osteichthyes: Sarcopterygii). Zoological Journal of the Lin-
nean Society 96, 119–166.
Ahlberg, P.E., 1992. A new holoptychiid porolepiform fish from the Upper
Frasnian of Elgin, Scotland. Palaeontology 35, 813–828.
Alekseev, A.S., Lebedev, O.A., Barskov, I.S., Barskova, M.I., Kononova, L.I.,
Chizhova, V.A., 1994. On the stratigraphic position of the Famennian and
Tournaisian fossil vertebrate beds in Andreyevka, Tula Region, Central Rus-
sia. Proceedings of the Geologist’s Association 105, 41–52.
Alekseev, A.S., Kononova, L.I., Nikishin, A.M., 1996. The Devonian and
Carboniferous of the Moscow Syneclise (Russian platform) and sea-level
changes. Tectonophysics 268, 149–168.
Anderson, M.E., Hiller, N., Gess, R.W., 1994. The first Bothriolepis-associated
Devonian fish fauna from Africa. South African Journal of Science 90,
397–403.
Anderson, M.E., Long, J.A., Evans, F.J., Almond, J.E., Theron, J.N., Bender,
P.A., 1999. Biogeographical affinities of Middle and Late Devonian fishes
of South Africa. Records of the Western Australia Museum 57, 157–168.
Andrews, S.M., 1978. A possible occurence of Remigolepis in the topmost
Old Red Sandstone of Berwickshire. Scottish Journal of Geology 14,
311–315.
Andrews, S.M., Long, J.A., Ahlberg, P., Barwick, R., Campbell, K., 2006. The
structure of the sarcopterygian fish Onychodus jandemarrai n. sp. from
Gogo, Western Australia: with a functional interpretation of the skeleton.
Transactions of the Royal Society of Edinburgh, Earth Sciences 96, 197–307.
Beznosov, P., 2002. Diversity and distribution of acanthodian remains in the
Upper Devonian of the South Timan (Russia). In: Brock, G.A., Talent,
J.A. (Eds.), First International Palaeontological Congress. 6–10 July, 2002,
Macquairie University, N.S.W., Australia. Geological Society of Australia,
Abstracts 68, p. 188.
Beznosov, P.A., 2009. Sosnogorskaya svita - novoye mestnoye stratigrafich-
eskoye podrazdeleniye verkhnego devona na Yuzhnom Timane [Sosnogorsk
Formation: a new local stratigraphic unit of the Upper Devonian in the South
Timan]. Geologiya i mineral’niye resursy severo-vostoka Evropeyskoy
Rossii. Materialy XV Geologicheskogo syezda Respubliki Komi [Geology
and Mineral Resources of the European North-East of Russia. Proceedings
of the XV Geological Congress of the Komi Republic], v. II. Syktyvkar, pp.
9–12 (in Russian).
Beznosov, P.A., Khipeli, D.V., Kuz’min, A.V., Belyayev, A.A., Khipeli, R.V.,
2004. Litologiya, ostatki pozvonochnykh i konodonty izhemskoy svity v
stratotipe [Lithology, vertebrate remains and conodonts of the Izhma For-
mation in the stratotype]. Geologiya i mineral’niye resursy severo-vostoka
Evropeyskoy Rossii. Materialy XIII syezda geologov Respubliki Komi
[Geology and Mineral Resources of the European North-East of Russia.
Proceedings of the XIII Geological Congress of the Komi Republic], vol.
III, Syktyvkar, pp. 220–224 (in Russian).
Blieck, A., Janvier, Ph., Lelièvre, H., Mistiaen, B., Montenat, Ch., 1982.
Vertébrés du Dévonien supérieur d’Afghanistan. Bulletin du Muséum
national d’Histoire Naturelle, Paris 4e sér., Section C 4, 3–19.
Blieck, A., Karatajute-Talimaa, V.N., Mark-Kurik, E., 2002. Upper Silurian
and Devonian heterostracan pteraspidomorphs (Vertebrata) from Severnaya
Zemlya (Russia): a preliminary report with biogeographical and biostrati-
graphical implications. Geodiversitas 24, 805–820.
Blom, H., Clack, J.A., Ahlberg, P.E., Friedman, M., 2007. Devonian verte-
brates from East Greenland: a review of faunal composition and distribution.
Geodiversitas 29, 119–141.
Clément, G., 2002. Large Tristichopteridae (Sarcopterygii, Tetrapodomorpha)
from the late Famennian Evieux Formation of Belgium. Palaeontology 45,
577–593.
Cloutier, R., Candilier, A.M., 1995. Palaeozoic vertebrates of northern France
and Belgium: Part III. Sarcopterygii (Devonian to Carboniferous). In:
Lelièvre, H., Wenz, S., Blieck, A., Cloutier, R. (Eds.), Premiers vertébrés et
vertébrés inferiéurs. Geobios, Mémoire spécial 19, pp. 335–341.
Daeschler, E.B., Frumes, A.C., Mullison, C.F., 2003. Groenlandaspid placoderm
fishes from the Late Devonian of North America. Records of the Australian
Museum 55, 45–60.
Denison, R., 1978. Placodermi. In: Handbook of Paleoichthyology. Gustav Fis-
cher Verlag, Stuttgart, New York, 128 pp.
Denison, R., 1979. Acanthodii. In: Handbook of Paleoichthyology. Gustav Fis-
cher Verlag, Stuttgart, New York, 62 pp.
Derycke, C., 1992. Microrestes de sélachiens et autres vertébrés du Dévonien
supérieur du Maroc. Bulletin du Muséum national d’Histoire Naturelle, Paris
4e sér., Section C 14, 15–61.
Eastman, C.R., 1907. Devonic fishes of the New York formations. Annual
Reports of the New York State Museum 60 (10), 1–235.
Elliott, D.K., Dineley, D.L., Johnson, H.G., 2000. A vertebrate fauna from the
Middle Devonian Yahatinda Formation of south-western Canada. Journal of
Paleontology 74, 123–132.
Elliott, D.K., Mark-Kurik, E., Daeschler, E., 2003. A revision of the obrucheviids
(Psammosteidae, Heterostraci) from the Late Devonian of the Arctic Canada
and Russia. In: Schultze, H.P., Luksevics, E., Unwin, D. (Eds.), The Gross
Symposium 2: Advances in Palaeoichthyology. Ichthyolith Issues, Special
Publication 7, pp. 21–22.
Esin, D., Ginter, M., Ivanov, A., Lebedev, O., Luksevics, E., Avkhi-
movich, V., Golubtsov, V., Petukhova, L., 2000. Vertebrate correlation
of the Upper Devonian and Lower Carboniferous on the East Euro-
pean Platform. In: Blieck, A., Turner, S. (Eds.), Palaeozoic Vertebrate
Biochronology and Global Marine-Non-Marine Correlation. Final Report
of IGCP 328 (1991–1996). Courier Forschungsinstitut Senckenberg 223,
pp. 341–359.
Friedman, M., Daeschler, E.B., 2006. Late Devonian (Famennian) lungfishes
from the Catskill Formation of Pennsylvania, USA. Palaeontology 49,
1167–1183.
Gardiner, B.G., 1984. The relationships of the palaeoniscid fishes, a review based
on new specimens of Mimia and Moythomasia from the Upper Devonian
of Western Australia. Bulletin of the British Museum (Natural History),
Geology 37, 173–428.
Ginter, M., 1999. Famennian-Tournaisian chondrichthyan microremains from
the Eastern Thuringian Slate mountains. Abhandlungen und Berichte für
Naturkunde 21, 25–47.
Ginter, M., 2000. Late Famennian pelagic shark assemblages. Acta Geologica
Polonica 50, 369–386.
Ginter, M., 2001. Chondrichthyan biofacies in the Late Permian of Utah and
Nevada. Journal of Vertebrate Paleontology 21, 714–729.
Ginter, M., 2002. Chondrichthyan fauna of the Frasnian-Famennian boundary
beds in Poland. Acta Palaeontologica Polonica 47, 329–338.
Ginter, M., Ivanov, A., 1992. Devonian phoebodont shark teeth. Acta Palaeon-
tologica Polonica 37, 55–75.
Ginter, M., Ivanov, A., 2000. Stratigraphic distribution of chondrichthyans
in the Devonian of the East European Platform margin. In: Blieck, A.,
Turner, S. (Eds.), Palaeozoic Vertebrate Biochronology and Global Marine-
Non-Marine Correlation. Final Report of IGCP 328 (1991–1996). Courier
Forschungsinstitut Senckenberg 223, pp. 325–339.
Ginter, M., Turner, S., 1999. The early Famennian recovery of phoebodont
sharks. Acta Geologica Polonica 49, 105–117.
Ginter, M., Hairapetian, V., Klug, Ch., 2002. Famennian chondrichthyans from
the shelves of North Gondwana. Acta Geologica Polonica 52, 169–215.
Greiner, H., 1977. Crossopterygian fauna from the Albert Formation, New
Brunswick, Canada, and its stratigraphic-paleoecologic significance. Journal
of Palaeontology 51, 44–56.
Gross, W., 1933. Die Fische des Baltischen Devons. Palaeontographica A 79,
1–74.
Gross, W., 1941. Die Bothriolepis-Arten der Cellulosa-Mergel Lettlands. Kung-
liga Svenska Vetenskaps-Akademiens Handlingar 19, 1–79.
Gross, W., 1942. Die Fischfaunen des baltischen Devons und ihre biostrati-
graphische Bedeutung. Korrespondentz-Blatt Naturforschunger Verein zu
Rı¯ga 64, 373–436.
Gross, W., 1950. Die paläontologische und stratigraphische Bedeutung der
Wirbeltierfaunen des Old Reds und der marinen altpaläozoischen Schichten.
Abhandlungen der Akademie der Wissenschaften zu Berlin 1, 1–130.
Gross, W., 1956. Über Crossopterygier und Dipnoer aus dem baltischen
Oberdevon im Zusammenhang einer vergleichenden Untersuchung des
Porenkanalsystems paläozoischer Agnathen und Fische. Kungliga Svenska
Vetenskaps-Akademiens Handlingar 5, 3–140.
Author's personal copy
126 O.A. Lebedev et al. / Palaeoworld 19 (2010) 108–128
Gross, W., 1973. Kleinschuppen. Flossenstacheln und Zähne von Fischen aus
Europäischen und nordamerikanischen Bonebeds des Devons. Palaeonto-
graphica A 142, 51–155.
House, M.R., Menner, V.V., Becker, R.T., Klapper, G., Ovnatanova, N.S.,
Kuzmin, A.V., 2000. Reef episodes, anoxia and sea-level changes in the
Frasnian of the Southern Timan. In: Insalaco, E., Skelton, P.W., Palmer, T.J.
(Eds.), Carbonate Platform Systems: Components and Interactions. Geolog-
ical Society of London. Special Publication 178, pp. 147–176.
Ivanov, A.O., 1990. Snetogorskiy kompleks ikhtiofauny Glavnogo devonskogo
polya i ego biostratigraficheskoye znacheniye [The Snetnaya Gora ichthy-
ofauna assemblage of the Main Devonian Field and its biostratigraphic
bearing]. Vestnik Leningradskogo Universiteta, Geologiya, Geografiya 7,
94–98 (in Russian).
Ivanov, A.O., 1999. Late Devonian-Early Permian chondrichthyans of the Rus-
sian Arctic. Acta Geologica Polonica 49, 267–285.
Ivanov, A.O., Ginter, M., 1997. Comments on the Late Devonian placoderms
from the Holy Cross Mountains (Poland). Acta Palaeontologica Polonica
42, 413–426.
Ivanov, A.O., Luksˇevicˇs, E., 1994. Famennian chondrichthyans from the East
European Platform. Daba un muzejs 5, 24–29.
Ivanov, A.O., Rodina, O., 2004. A new omalodontid-like shark from the Late
Devonian (Famennian) of western Siberia, Russia. Fossils and Strata 50,
82–91.
Ivanov, A.O., Esin, D.N., Vyushkova, L.V., 1992. Ichthyofauna. In: Kras-
nov, V.I., Rzhonsnitskaya, M.A., Gutak, Ya.M. (Eds.), Tipoviye razrezy
pogranichnykh otlozheniy srednego i verkhnego devona, frana i famena
okrain Kuznetskogo basseyna [The Type Sections of the Boundary Deposits
of the Middle and Upper Devonian, Frasnian and Famennian in the Periphery
of the Kuznetsk Basin]. Materialy V sessii devonskoy mezhvedomstvennoy
stratigraficheskoy komissii [Materials of the V Session of the Interdepart-
mental Stratigraphical Commission on the Devonian], Novosibirsk, pp.
89–91 (in Russian).
Ivanov, A.O., Luksˇevicˇs, E., Stinkulis, G¸., Tovmasjan, K., Zupin¸sˇ, I., Beznosov,
P.A., 2006. Stratigrafiya devona Andomskoy gory [Stratigraphy of the Devo-
nian at the Andoma Hill]. In: Pystin, A.M. (Ed.), Problemy geologii i
mineralogii. Syktyvkar, pp. 385–396 (in Russian).
Janvier, P., 1983. Les vertébrés dévoniens de la Nappe Superieure d’Antalya
(Taurus Lycien occidental, Turquie). Géologie Méditerranee 10, 1–13.
Janvier, Ph., 2003. The Devonian vertebrates from Bolivia: new data, more enig-
mas. Revista Técnica de Yacimientos Petrolíferos Fiscales Bolivianos 21,
25–35.
Janvier, P., Martin, M., 1979. Les vertebres devoniens de l’Iran central. II.
Coelacanthiformes, Struniiformes, Osteolepiformes. Geobios 12, 497–511.
Janvier, P., Villarroel, C., 2000. Devonian vertebrates from Colombia. Palaeon-
tology 43, 729–763.
Janvier, P., Clément, G., Cloutier, R., 2007. A primitive megalichthyid fish (Sar-
copterygii, Tetrapodomorpha) from the Upper Devonian of Turkey and its
biogeographical implications. Geodiversitas 29, 249–268.
Jarvik, E., 1950. Note on Middle Devonian crossopterygians from the eastern
part of Gauss Halvo, East Greenland. With an appendix: an attempt at a
correlation of the Upper Old Red Sandstone. Meddelelser om Grønland
149, 3–20.
Jarvik, E., 1972. Middle and Upper Devonian Porolepiformes from East Green-
land with special references to Glyptolepis groenlandica n. sp. Meddelelser
om Grønland 187, 1–295.
Jessen, H., 1966. Die crossopterygier des oberen Plattenkalkes (Devon) der
Bergisch-Gladbach-Paffrather Mulde (Rheinisches Schiefergebirge) unter
Berücksichtigung von amerikanischem und europäischem Onychodus-
Material. Arkiv för zoologi 18, 305–391.
Johanson, Z., 1997. New Remigolepis (Placodermi; Antiarchi) from
Canowindra, New South Wales, Australia. Geological Magazine 134,
813–846.
Johanson, Z., Ritchie, A., 2000. Rhipidistians (Sarcopterygii) from the Hunter
Siltstone (Late Famennian) near Grenfell, NSW, Australia. Mitteilungen aus
dem Museum für Naturkunde in Berlin 3, 111–136.
Krupina, N.I., 1986. O kryshe cherepa Orlovichthys limnatis (Dipnoi) [On the
skull roof of Orlovichthys limnatis (Dipnoi)]. Paleontologicheskiy Zhurnal
2, 104–108 (in Russian).
Krupina, N.I., 1987. Novaya dvoyakodyshashchaya ryba iz verkhnego devona
Tul’skoy oblasti [A new dipnoan from the Upper Devonian of the Tula
Region]. Paleontologicheskiy Zhurnal 3, 40–47 (in Russian).
Krupina, N.I., 1999. Novaya dvoyakodyshashchaya ryba iz verkhnedevonskogo
mestonakhozhdeniya Rybnitsa Orlovskoy oblasti [A new dipnoan from the
Upper Devonian locality Rybnitsa of the Oryol Region]. Paleontologicheskiy
Zhurnal 6, 24–26 (in Russian).
Krupina, N.I., 2000. Dvoyakodyshashchiye ryby iz verkhnedevonskogo meston-
akhozhdeniya Rybnitsa (Orlovskaya oblast’) [Dipnoans from the Upper
Devonian locality Rybnitsa (Oryol Region)]. Paleontologicheskiy Zhurnal
5, 55–61 (in Russian).
Krupina, N.I., 2004. Podklass Dipnoi. Dvoyakodyshashchiye. [Subclass Dipnoi.
Dipnoans]. In: Novitskaya, L.I., Afanassieva, O.B. (Eds.), Iskopayemiye
pozvonochniye Rossii i sopredel’nykh stran. Beschelyustniye i drevniye
ryby [Fossil Vertebrates of Russia and Adjacent Countries. Agnathans and
Early Fishes]. Geos Publishers, Moscow, pp. 373–413 (in Russian).
Krupina, N.I., Reisz, R.R., 1999. Reconstruction of dentition in hatchlings of
Andreyevichthys epitomus, a late Famennian dipnoan from Russia. Modern
Geology 24, 99–108.
Krylova, A.K., 1973. Donetskiy basseyn [The Donets Basin]. In: Nalivkin, D.V.,
Rzonsnitskaya, M.A., Markovskiy, B.P. (Eds.), Stratigrafiya SSSR. Devon-
skaya sistema [Stratigraphy of the USSR. The Devonian System], vol. 1.
Nedra Publishers, Moscow, pp. 201–206 (in Russian).
Kulczycki, J., 1957. Upper Devonian fishes from the Holy Cross Mountains
(Poland). Acta Palaeontologica Polonica 2, 285–380.
Lebedev, O.A., 1983. Novaya kisteperaya ryba s territorii Tsentral’nogo devon-
skogo polya [A new crossopterygian fish from the territory of the Central
Devonian field]. Paleontologicheskiy Zhurnal 4, 68–75 (in Russian).
Lebedev, O.A., 1985. Perviye tetrapody: poiski i nakhodki [The first tetrapods:
searches and finds]. Pririda 11, 26–36 (in Russian).
Lebedev, O.A., 1992. The latest Devonian, Khovanian vertebrate community of
Andreyevka-2 locality, Tula Region, Russia. In: Mark-Kurik, E. (Ed.), Fossil
fishes as living animals. Proceedings of the II International Colloquium on
the Study of the Palaeozoic fishes, Tallinn, 1989. Academia, vol. 1, pp.
265–272.
Lebedev, O.A., 1995. Middle Famennian (Upper Devonian) chondrichthyans
and sarcopterygians from Oryol Region, Central Russia. In: Lelièvre, H.,
Wenz, S., Blieck, A., Cloutier, R. (Eds.), Premiers vertébrés et vertébrés
inferiéurs. Geobios, Mémoire spécial 19, pp. 361–368.
Lebedev, O.A., 2004. A new tetrapod Jakubsonia livnensis from the early
Famennian (Devonian) of Russia and palaeoecological remarks on the Late
Devonian tetrapod habitats. Acta Universitatis Latviensis, Earth and Envi-
ronment Science, Special Volume 679, 79–98.
Lebedev, O.A., Luksˇevicˇs, E., 1996. Attempted correlation of the upper part
of the Famennian deposits of Baltic and Central Russia by vertebrates. In:
Meidla, T., Puura, I., Nemliher, J., Raukas, A., Saarse, L. (Eds.), The Third
Baltic Stratigraphical Conference. Abstracts, Field Guide. Tartu, pp. 35–36.
Lebedev, O.A., Zakharenko, G.V., 2010. Global vertebrate-based Palaeozoo-
geographical regionalisation during the Givetian-Famennian (Middle-Late
Devonian). Endemism-cosmopolitanism spectrum as an indicator of inter-
provincial faunal exchanges. Palaeoworld 19 (1–2), 186–205.
Lelièvre, H., Janvier, Ph., Blieck, A., 1993. Silurian-Devonian vertebrate bios-
tratigraphy of Western Gondwana and related terranes (South America,
Africa, Armorica-Bohemia, Middle East). In: Long, J.A. (Ed.), Palaeozoic
Vetrebrate Biostratigraphy and Biogeography. Belhaven Press, London, pp.
139–173.
Liszkowski, J., Racki, G., 1992. Ichthyoliths and deepening events in the Devo-
nian carbonate platform of the Holy Cross Mountains. Acta Palaeontologica
Polonica 37, 407–426.
Long, J.A., 1993. Early-Middle Palaeozoic vertebrate extinction events. In:
Long, J.A. (Ed.), Palaeozoic Vetrebrate Biostratigraphy and Biogeography.
Belhaven Press, London, pp. 54–63.
Long, J.A., Hairapetian, V., 2000. Famennian microvertebrates from the
Dalmeh area, central Iran. Records of the Western Australia Museum 58,
211–221.
Luksˇevicˇs, E., 1991. Finding of Remigolepis (Pisces, Antiarchi) in the Famen-
nian deposits of the Central Devonian Field (Russia, Tula government). Daba
un muzejs 3, pp. 51–56 (in Russian, with Latvian and English summaries).
Author's personal copy
O.A. Lebedev et al. / Palaeoworld 19 (2010) 108–128 127
Luksˇevicˇs, E., 1999. Stratigraphical occurence of vertebrate remains in the
Upper Devonian of Severnaya Zemlya (Russia). Acta Geologica Polonica
49, 125–131.
Luksˇevicˇs, E., 2001. Bothriolepid antiarchs (Vertebrata, Placodermi) from the
Devonian of the north-western part of the East European Platform. Geodi-
versitas 23, 489–609.
Luksˇevicˇs, E., Zupin¸sˇ, I., 2004. Sedimentology, fauna, and taphonomy of the
Pava¯ri site, Late Devonian of Latvia. Acta Universitatis Latviensis, Earth
and Environmental Sciences 679, 99–119.
Luksˇevicˇs, E., Lebedev, O.A., Zakharenko, G.V., 2010. Palaeozoogeographical
connections of the Devonian vertebrate communities of the Baltica Province.
Part I. Early-Middle Devonian. Palaeoworld 19 (1–2), 108–128.
Malinovskaya, S., 1992. New Middle Devonian antiarchs (Placodermi) of Cen-
tral Kazakhstan. In: Mark-Kurik, E. (Ed.), Fossil Fishes as Living Animals.
Proceedings of the II International Colloquium on the Study of the Palaeozoic
fishes, Tallinn, 1989. Academia, vol. 1, pp. 177–184.
Mark-Kurik, E., 1974. Discovery of a new Devonian fish localities in the Soviet
Arctic. Eesti NSV Teaduste Akadeemia Toimetised 23, 332–335.
Matukhin, R.G., Menner, V.V., 1999. Stratigrafiya silura i devona arkhipelaga
Severnaya Zemlya [Stratigraphy of the Silurian and Devonian of the Sever-
naya Zemlya Archipelago]. SNIIGGiMS, Novosibirsk, 174 pp. (in Russian).
Miles, R.S., 1968. The Old Red Sandstone antiarchs of Scotland: Family Both-
riolepididae. Palaeontographical Society Monographs 522, 3–130.
Miles, R.S., 1977. Dipnoan (lungfish) skulls and the relationships of the group:
a study based on new species from the Devonian of Australia. Zoological
Journal of the Linnean Society 61, 1–328.
Moloshnikov, S.V., 2001. Unikal’noye mestonakhozhdeniye pozvonochnykh
verkhnego devona Orlovskoy oblasti [A unique vertebrate locality in the
Upper Devonian of the Oryol Region]. Izvestiya VUZov. Geologiya i
Razvedka 3, 29–33 (in Russian).
Moloshnikov, S.V., 2002. Franskiye antiarkhi (Pisces, Placodermi) Tsen-
tral’nogo devonskogo polya [Frasnian antiarchs of the Central Devonian
field]. Izvestiya VUZov, Geologiya i Razvedka 4, 12–19, 159 (in Russian).
Moloshnikov, S.V., 2004. Crested antiarch Bothriolepis zadonica H.D.
Obrucheva from the Lower Famennian of Central European Russia. Acta
Palaeontologica Polonica 49, 135–146.
Moloshnikov, S.V., 2007. O nakhodkakh psammosteid (Agnatha, Heterostraci) v
nizhnefranskikh otlozheniyakh Mikhailovskogo rudnika (Kurskaya oblast’)
[On the psammosteid finds in the Lower Frasnian deposits of the Mikhailov
quarry (Kursk Region)]. Paleontologicheskiy Zhurnal 5, 84–88 (in Russian).
Moloshnikov, S.V., 2008. Devonian antiarchs (Pisces, Antiarchi) from
Central and Southern European Russia. Paleontological Journal 42,
691–773.
Newberry, J.S., 1889. The Paleozoic fishes of North America. US Geological
Survey Monographs 16, 1–228.
Obruchev, D.V., 1940. Devonskiye ryby Sibiri I Sredney Azii [The Devonian
fishes of Siberia and Central Asia]. Doklady Akademii Nauk SSSR 27,
889–892 (in Russian).
Obruchev, D.V., 1941. Devonskiye ryby Minusinskogo kraya [The Devonian
fishes of Minusa]. Trudy PIN AN SSSR 8, 23–48 (in Russian).
Obruchev, D.V., 1955. Devonskiye ryby Minusinskoy kotloviny [The Devonian
fishes of the Minusa depression]. In: Rzhonsnitskaya, M.A., Meleshchenko,
V.S. (Eds.), Polevoy atlas kharakternykh kompleksov fauny i flory
devonskikh otlozheniy Minusinskoy kotloviny [The Field Atlas of the Char-
acteristic Assemblages of Fauna and Flora in the Devonian deposits of the
Minusa depression]. VSEGEI, Leningrad, pp. 45–47 (in Russian).
Obruchev, D.V., 1958. O biostratigrafii nizne- i srednepaleozoyskikh ikhtio-
faun SSSR [On the biostratigraphy of the Lower and Middle Palaeozoic
ichthyofaunas of the USSR]. Sovetskaya geologiya 1, 40–53 (in Russian).
Obruchev, D.V., 1959. Noviy vid Palaedaphus (Dipnoi) iz verkhnego devona
Russkoy platformy [A new species of Palaedaphus (Dipnoi) from the Upper
Devonian of the Russian platform]. Paleontologicheskiy Zhurnal 4, 145–146
(in Russian).
Obruchev, D.V., 1960. Ikchtiofauna [Ichthyofauna]. In: Khalfin, L.L. (Ed.), Bios-
tratigrafiya paleozoya Sayano-Altayskoy skladchatoy oblasti [Palaeozoic
Biostratigraphy of the Sayan-Altay Folded Belt], v. 2. Trudy SNIIGGiMS
20, pp. 296–301 (in Russian).
Obruchev, D.V., Sergienko, A.A., 1960. Placodermi. Plastinokozhiye [Pla-
codermi. Placoderms]. In: Khalfin, L.L. (Ed.), Biostratigrafiya paleozoya
Sayano-Altayskoy skladchatoy oblasti [Palaeozoic Biostratigraphy of the
Sayan-Altay Folded Belt], v. 2. Trudy SNIIGGiMS 20, pp. 561–564 (in
Russian).
Obrucheva, E.D., 1983. A new bothriolepid species from the deposits of
the Zadonsk Member of the Central Devonian Field. In: Novitskaya,
L.I. (Ed.), Problemy sovremennoy paleoikhtiologii (Materialy konferentsii,
posvyashchennoy D.V. Obruchevu) [Problems of Modern Palaeoikhthyol-
ogy (Materials of the D.V. Obruchev Conference)]. Nauka, Moscow, pp.
36–42 (in Russian).
Obrucheva, O.P., 1956. Ostatki Dinichthys (Arthrodira) iz verkhnego devona
SSSR [Remains of Dinichthys (Arthrodira) from the Upper Devo-
nian of the USSR]. Doklady Akademii Nauk SSSR 108, 333–336
(in Russian).
Obrucheva, O.P., 1962. Pantsirniye ryby devona SSSR (kokkosteidy i dinikhtidy)
[Armoured fishes from the Devonian of USSR (coccosteids, dinichthyids)].
MGU Publishing House, Moscow, 189 pp. (in Russian).
Obrucheva, O.P., 1983. Rod Chelyophorus iz otlozheniy Tsentral’nogo devon-
skogo polya [The genus Chelyophorus from the deposits of the Central
Devonian field]. In: Novitskaya, L.I. (Ed.), Problemy sovremennoy pale-
oikhtiologii (Materialy konferentsii, posvyashchennoy D.V. Obruchevu)
[Problems of Modern Palaeoikhthyology (Materials of the D.V. Obruchev
Conference)]. Nauka, Moscow, pp. 28–36 (in Russian).
Obrucheva, O.P., Obrucheva, E.D., 1977. Fishes of the Central Devonian field.
In: Menner, V.V. (Ed.), Ocherki po filogenii I sistematike iskopayemykh ryb
i beschelyustnykh [Essays on the Phylogeny and Systematics of the Fossil
Fishes and Agnathans]. Nauka, Moscow, pp. 24–28 (in Russian).
Ørvig, T., 1957. Remarks on the vertebrate fauna of the Lower Upper Devonian
of Escuminac Bay, P.Q., Canada, with special reference to the Porolepiform
Crossopterygians. Arkiv for Zoologi 10, 367–426.
Panteleyev, N., 1992. New remigolepids and high armoured antiarchs of Kirgizia.
In: Mark-Kurik, E. (Ed.), Fossil Fishes as Living Animals. Proceedings of
the II International Colloquium on the Study of the Palaeozoic fishes, Tallinn,
1989. Academia, vol. 1, pp. 185–191.
Randon, C., Derycke, C., Blieck, A., Perri, M.C., Spalletta, C., 2007. Late
Devonian–Early Carboniferous vertebrate microremains from the Carnic
Alps, northern Italy. Geobios 40, 809–826.
Rieman, F., Scholke, I., Thies, D., 2002. Mikrovertebratenreste aus dem basalen
Famennium (triangularis- bis crepida-Zone) der Montagne Noire (Frankre-
ich). Geologica et Palaeontologica 36, 1–43.
Rodionova, G.D., Umnova, V.T., Kononova, L.I., Ovnatanova, N.S., Rzhon-
snitskaya, M.A., Fedorova, T.I., 1995. Devon Voronezhskoy anteklizy i
Moskovskoy sineklizy [The Devonian of the Voronezh Anteclise and the
Moscow Syneclise]. Rosgeolfond, Moscow, 265 pp. (in Russian).
Savvaitova, L., 1977. Famen Pribaltiki. Zina¯tne, Riga, 128 pp. (in Russian).
Schultze, H.P., 1973. Large Upper Devonian arthrodires from Iran. Fieldiana:
Geology 23, 53–78.
Schultze, H.P., Cloutier, R., 1996. Comparison of the Escuminac Forma-
tion ichthyofauna with other Late Givetian/early Frasnian ichthyofaunas.
In: Schultze, H.P., Cloutier, R. (Eds.), Devonian Fishes and Plants
of Miguasha, Quebec, Canada. Verlag Dr. Friedrich Pfeil, München,
pp. 348–368.
Sorokin V.S., Lyarskaya, L.A., Savvaitova, L.S., 1981. Devon i karbon Pribaltiki
[Devonian and Carboniferous of the Peribaltic Region]. Zina¯tne, Riga, 502
pp. (in Russian).
Stinkulis, G¸., 2008. Dolocretes in the Devonian deposits of Latvia. In: Hints, O.,
Ainsaar, L., Mannik, P., Meidla, T. (Eds.), The Seventh Baltic Stratigraphi-
cal Conference. Abstracts and Field Guide. Geological Society of Estonia,
Tallinn, p. 66.
Tikhomirov, S.V., 1967. Etapy osadkonakopleniya devona Russkoy platformy
[Phases of deposition of the Devonian on the Russian Platform]. Nedra,
Moscow, 267 pp. (in Russian).
Tovmasyan, K., Stinkulis, G., 2008. Implications of fluvial, tidal and wave pro-
cesses to the deposition of siliciclastic succession of Devonian, Andoma
Hill, NW Russia. In: Hints, O., Ainsaar, L., Männik, P., Meidla, T. (Eds.),
The Seventh Baltic Stratigraphical Conference. Abstracts and Field Guide.
Geological Society of Estonia, Tallinn, p. 71.
Author's personal copy
128 O.A. Lebedev et al. / Palaeoworld 19 (2010) 108–128
Turner, S., Young, G.C., 1997. Devonian vertebrate faunal response to eustatic
sea-level changes in East Gondwana. In: House, M.R., Ziegler, W. (Eds.), On
Sea-Level Fluctuations in the Devonian. Courier Forschungsinstitut Senck-
enberg 199, pp. 117–127.
Valiukevicˇius, J., Kruchek, S., 2000. Acanthodian biostratigraphy and interre-
gional correlations of the Devonian in the Baltic States, Belarus, Ukraine
and Russia. In: Blieck, A., Turner, S. (Eds.), Palaeozoic Vertebrate
Biochronology and Global Marine-Non-Marine Correlation. Final Report
of IGCP 328 (1991–1996). Courier Forschungsinstitut Senckenberg 223,
pp. 271–289.
Vinogradov, A.P., Nalivkin, V.D. (Eds.), 1960. Atlas litologo-paleogeo-
graficheskikh kart Russkoy platformy i yeyo geosinklinal’nogo obramleniya.
[Atlas of the lithological-palaeogeographical maps of the Russian platform
and its geosynclinal rim]. Part 1. Late Precambrian and Palaeozoic. Gosge-
olizdat, Moscow-Leningrad, maps 17–23.
Vorobyeva, E.I., 2004. Klass Sarcopterygii. Sarkopterigii. [Class Sarcoptery-
gii. The Sarcopterygians]. In: Novitskaya, L.I., Afanassieva, O.B. (Eds.),
Iskopayemiye pozvonochniye Rossii i sopredel’nykh stran. Beschelyust-
niye i drevniye ryby. [Fossil Vertebrates of Russia and Adjacent Countries.
Agnathans and Early Fishes]. Geos Publishers, Moscow, pp. 271–372 (in
Russian).
Vorobyeva, E.I., Obruchev, D.V., 1964. Podklass Sarcopterygii. In: Obruchev,
D.V. (Ed.), Fundamentals of Palaeontology. Agnathans, Fishes, Nauka,
Moscow, pp. 238–266 (in Russian).
Vorobyeva, E.I., Panteleev, N.V., Kolobayeva, O.V., Blieck, A., 1997. Upper
Devonian ichthyofauna from the Michailovskij excavation of the Kursk
magnetic anomaly, Russia: a preliminary report. In: Wimbledon, W.A.,
Fraser, N.C. (Eds.), Gross Symposium. Modern Geology Special Issue 21,
79–86.
Young, G.C., 1988. Antiarchs (placoderm fishes) from the Devonian Aztec
Siltstone, Southern Victoria Land, Antarctica. Palaeontographica A 202,
1–125.
Young, G.C., 2003. North Gondwana mid-Palaeozoic connections with
Euramerica and Asia: Devonian vertebrate evidence. Courier Forschungs-
Institut Senckenberg 242, 169–185.
Young, G.C., 2006. Biostratigraphic and biogeographical context for tetrapod
origins during the Devonian: Australian evidence. Alcheringa Special Issue
1, 409–428.
Young, G.C., Long, J.A., 2005. Phyllolepid placoderm fish remains from the
Devonian Aztec Siltstone, southern Victoria Land, Antarctica. Antarctic
Science 17, 387–408.
Young, G.C., Turner, S., 2000. Devonian microvertebrates and
marine–nonmarine correlation in East Gondwana: overview. In: Blieck,
A., Turner, S. (Eds.), Palaeozoic Vertebrate Biochronology and Global
Marine-Non-Marine Correlation. Final Report of IGCP 328 (1991–1996).
Courier Forschungsinstitut Senckenberg 223, pp. 453–470.
Young, G.C., Moody, J.M., Casas, J.E., 2000. New discoveries of Devonian ver-
tebrates from South America, and implications for Gondwana-Euramerica
contact. Comptes Rendus Academie Sciences de Paris. Sciences de la terre
et de planetes 331, 755–761.
Zakharenko, G.V., 2007. Novaya pakhiosteomorfnaya artrodira (Pisces: Pla-
codermi) iz verkhnego frana Tsentral’nogo devonskogo polya Rossii [A
new pachyosteomorph arthrodire (Pisces: Placodermi) from the Upper Fras-
nian of the Central Devonian Field, Russia]. Paleontologicheskiy Zhurnal 6,
642–655 (in Russian).
Zangerl, R., 1981. Chondrichthyes I. Paleozoic Elasmobranchii. In: Handbook
of Paleoichthyology. Gustav Fischer Verlag, Stuttgart, New York, 115 pp.
Zhao, W.J., Zhu, M., 2007. Diversification and faunal shift of Siluro-Devonian
vertebrates of China. Geological Journal 41, 1–17.
Zhu, M., 2000. Catalogue of Devonian vertebrates in China, with notes
on bio-events. In: Blieck, A., Turner, S. (Eds.), Palaeozoic Vertebrate
Biochronology and Global Marine-Non-Marine Correlation. Final Report
of IGCP 328 (1991–1996). Courier Forschungsinstitut Senckenberg 223,
pp. 373–390.