ISSN 08695938, Stratigraphy and Geological Correlation, 2012, Vol. 20, No. 2, pp. 211229. Pleiades Publishing, Ltd.

., 2012.
Original Russian Text M.A. Rogov, D.B. Gulyaev, D.N. Kiselev, 2012, published in Stratigrafiya. Geologicheskaya Korrelyatsiya, 2012, Vol. 20, No. 2, pp. 101121.

Biohorizons as Infrazonal Biostratigraphic Units: An Attempt

to Refine the Jurassic Stratigraphy Based on Ammonites
M. A. Rogova, D. B. Gulyaevb, and D. N. Kiselevc
aGeological
Institute, Russian Academy of Sciences, Pyzhevskii per. 7, Moscow, 119017 Russia
email: russianurassic@gmail.com
b
ScientificIndustrial Center Nedra, ul. Svobody 8/38, Yaroslavl, 150000 Russia
email: dgulyaev@rambler.ru
c
Yaroslavl Ushinskii State Pedagogical University, ul. Respublikanskaya 108, Yaroslavl, 150000 Russia
email: dnkiselev@mail.ru
Received February 8, 2011; in final form, June 7, 2011

AbstractThe biohorizons (faunal horizons) as infrazonal units are the smallest correlatable biostratigraphic
units. Their main features are: (1) potential indivisibility based on taxonomic differentiation of guide fossils;
(2) determinancy of both lower and upper boundaries in the geological section; (3) identification by a single
index species/subspecies. First such units were defined at the end of the 19th century and since the 1980s have
been widely used in biostratigraphic investigations of Jurassic and, later, Cretaceous systems. The biohorizons
are characterized by phylogenetic or immigrational paleobiological nature and geologically they are con
nected with depositional and postdepositional transformation (and, consequently, structure) of the sedimen
tary succession. Based on parallel sequences of phylogenetic and separate immigrational biohorizons, they
are integrated into different zonal scales and an integrated regional scale. The problems related to the lack of
universal criteria for defining and using biohorizons are discussed. The basic nomenclature rules, which are
aimed at regulation of the use of these units in practical stratigraphic investigations, are suggested for their
recognition and description.

Keywords: infrazonal biostratigraphy, biohorizons, faunal horizons, ammonites.

As the middle system of the middle

(Mesozoic) group of fossiliferous rocks,
the Jurassic is as good a sample as any
that could be chosen to find out what
light can be thrown on geological prob
lems by the study of a single system all
over the world (Arkell, 1956, p. 3).
DOI: 10.1134/S0869593812010066

INTRODUCTION
Despite the 150yearlong development history of
the zonal method in stratigraphy, its basic theoretical
principles concerning biostratigraphic zones were for
mulated only in the second half of the 20th century. At
the end of the 19th century, it became clear that
smaller (infrazonal) biostratigraphic units may also be
useful for subdivision and correlation of sedimentary
sections.
Now, these smallest infrazonal biostratigraphic
units (biohorizons, faunal horizons) are widely used in
biostratigraphic investigations of the Jurassic and Cre
1
taceous systems. Similar to many other innovations
in stratigraphy, paleobiology, and paleogeography, the
infrazonal biostratigraphic units were first proposed by
the experts engaged in study of the Jurassic System and
are now most widely used in the Jurassic stratigraphy.
Until recently, the biohorizons (in the sense consid
ered in this work) are defined by stratigraphers only for
1 Some
researchers (Page, 1995; and others) use in this sense the
term intrasubzonal units. Nevertheless, inasmuch as the sub
zones represent stratigraphic units of the zonal rank (similar to
subgenera, which are taxa of the generic rank), the use of the less
cumbersome adjective infrazonal seems a better choice for
biohorizons.

211
212 ROGOV et al.

enodatum planicerclus

Sigaloceras calloiense
Catasigaloceras pagei
enodatum enodatum
(after Kiselev, 2001;

Kepplerites galilaeii
Gulyaev et al.2002)

Sigaloceras micans
enodatum aeeta
Catasigaloceras

Catasigaloceras

Catasigaloceras
medea medea
Kosmoceras
Biohorizons

Spinosum

Grossouvrei

Gowerianus
Calloviense
Obductum

Terebratus
Enodatum

Curtilobus
Phaeinum
Lamberti

Kamptus

Keppleri
Galilaeii
Proniae
Henrici
Substages, zone, and subzones

Medea
Jason
Coronatum

Calloviense
Lamberti

Koenigi

Herveyi
Athleta

Jason

Upper Middle Lower

Kimmeridgian

Pliensbachian

Sinemurian

Hettangian
Series and stages

Oxfordian

Bathonian
Tithonian

Callovian

Aalenian

Toarcian
Bajocian

Upper Middle Lower

Erathems and systems

Cretaceous

Jurassic

Triassic

Paleozoic
Cenozoic

Mesozoic

Fig. 1. The growth of resolution of stratigraphic scales depending on the lower rank of stratigraphic units.

the Mesozoic based mainly on ammonites, although it possible to increase the biostratigraphic resolution
the number of fossil groups used for defining such several times (Fig. 1) and the accuracy of correlation
stratigraphic units as well as for widening the strati between biostratigraphic scales (Fig. 2).
graphic range of their application is gradually increas At the same time, the use of infrazonal biostrati
ing. The application of the infrazonal approach made graphic units is frequently ambiguous and even slightly

STRATIGRAPHY AND GEOLOGICAL CORRELATION Vol. 20 No. 2 2012

 BIOHORIZONS AS INFRAZONAL BIOSTRATIGRAPHIC UNITS 213

Stage Biochoremas
Boreal Atlantic realm East MediterraneanCaucasian Realm
European province Submediterranean province
(European Russia) (mainly central and southern France, Portugal)
(Fig. 7) (after Biostratigraphie , 1997)

Upper (pars) Substage

Substage

Biohorizons

Zone
Zone
Subzone (tints show biohorizons of Biohorizons Subzone
different subprovinces)
Collotia collotiformis
Spinosum K. kuklikum Collotiformis

Athleta
Orionoides piveteaui
Upper (pars)
Athleta

F. funiferus
K. rowlstonense Sublunuloceras trezeense
Proniae F. patruus Trezeense
F. sp. nov. K. proniae Pseudopeltoceras leckenbyi
Pseudopeltoceras Rota
Phaeinum K. phaeinum
F. allae Rehmannia rota

Coronatum
Coronatum

K. grossouvrei Choffatia waageni

Grossouvrei Leuthardti
K. posterior Erymnoceras leuthardti
Callovian

K. crassum Erymnoceras baylei

Middle
Obductum L. stenolobum Baylei
Middle

L. praestenolobum K. obductum Flabellisphinctes villanyensis

K. jason jason Rehmannia richei
Jason Tyranniformis

Anceps
K. jason
Jason

sedgwickii Rehmannia blyensis

K. medea magnum Chanasia turgida

Medea R. milaschevici Stuebeli
milaschevici
K. medea medea Chanasia bannensis
?

Gracilis (pars)
Lower (pars)
C. enodatum aeeta
Lower (pars)
Calloviense

R. milaschevici C. enodatum Hecticoceras posterius

(pars)

Enodatum khudyaevi enodatum Patina

i A. difficilis C. enodatum Collotia pamprouxensis
R. fkin planicerclus
e f
tch P. cracoviensis C. pagei Hecticoceras boginense

Fig. 2. Correlation of biohorizons in regional (provincial) stratigraphic scales for the Middle Callovian in different paleobiogeo
graphic realms. Here and in Figs. 37, names of conditional (preliminary) biohorizons are given in quotes. The dark and light
gray colors indicate sequences of biohorizons based on cardioceratids and perisphinctids, respectively.

chaotic. In this work, we consider critically the devel zonal units, for example, the Ammonites calloviensis
opment history of the infrazonal biostratigraphy, ana and A. bullatus horizons in the A. macrocephalus Zone
lyze the main features and nature of biohorizons, (Oppel, 18561858, p. 2009). Slightly later, Waagen
assess their correlation potential, and formulate prin (1869), who established the first phylogenetic lineage
ciples of their recognition and application. In addi of Bathonian and Callovian ammonites, formulated a
tion, we propose a code of rules, which should concept of evolutionary mutations (quantitative mor
decrease, if not eliminate, ambiguity inherent now in phological transformations) through time and
the infrazonal biostratigraphy. described the zone as an interval corresponding to the
existence period of a single mutation. This concept
was further developed by Neumayr (1878, p. 40), who
A DEVELOPMENT HISTORY OF INFRAZONAL noted that the Oppels zones of the Jurassic System
BIOSTRATIGRAPHY chronologically correspond to the average existence
In his famous work dedicated to the Jurassic Sys period of a single mutation among most widespread
tem of England, France, and southwestern Germany, marine animals primarily such as cephalopods.
Oppel (18561858), who formulated the basics of the Already at that time, the inconsistency between the
zonal stratigraphy, described the zone as a horizon, historical continuity of zonal scales and their resolu
which is characterized by the same fossil assemblage tion became clear for many researchers of the Jurassic
even in remote areas of its development. He empha System. The zones that became generally accepted at
sized that the accuracy in defining zones depends on that time could be divided into smaller units. Thus,
the number of well described species (especially guide two different approaches to the zonal stratigraphy
taxa), in addition to other factors. In the same work, existed at the end of the 19th century. The zones were
this author defined the socalled horizons as intra accepted either as smallest biostratigraphic units or as

STRATIGRAPHY AND GEOLOGICAL CORRELATION Vol. 20 No. 2 2012

214 ROGOV et al.
some stable stratigraphic framework, which implied
the possibility of defining smaller stratigraphic units.
The further development of views on detailed
stratigraphy is primarily connected with S. Buckman
and his undeservedly forgotten concept of a hemera.
The latter was in fact understood as a geochronologi
cal equivalent of the biohorizon. The last researcher
defined it as a chronological indicator of the small
est stratigraphic interval corresponding to the acme of
single or several species (Buckman, 1893, 1902).
When studying limestones from the Inferior Oolite
Group in the Dorset and South Somerset (England),
Buckman (1893) was able to compile an unusually
detailed sequence of faunal assemblages based on
thorough bedbybed sampling of ammonites and cor
relation of their host sections. The matter was compli
cated by the fact that the Lower Oolite rocks are
strongly condensed and none of their sections is strati
graphically complete. Practically every bed with
ammonites is separated from its neighboring counter
parts by distinct or cryptic hiatuses. In order to over
come this difficulty, Buckman first established the
bedbybed sequence of ammonite assemblages in
each section. Then, this researcher correlated identi
cal assemblages from different sections with certain
time intervals (hemerae) indexing them with most
characteristic species or genera. He arranged the chro
nological succession of the hemerae correlating suc
cessions of characteristic assemblages in many sec
tions. The recent revision of Middle Jurassic sections
formerly studied by Buckman confirmed his strati
graphic interpretations (Callomon and Chandler,
1990). It should be emphasized that in the Buckmans
understanding, hemerae were precisely chronological,
not stratigraphic units representing time equivalents of
zones (Buckman, 1902). Because of the ambiguity in
the use of the term zone in different Buckmans
works, Trueman (1923) suggested to use an epibole
as a stratigraphic equivalent of the hemera.
In his subsequent works, Buckman (19091930)
extended his method practically to the entire Jurassic
System of England. He united hemerae into ages
largely corresponding to the stratigraphic ranges of
particular ammonite families or genera. Buckman
considered the composite sequence of hemerae and
ages as a geochronological calendar. He emphasized
that the hemera corresponds to the existence period of
the index species (Buckman, 1898). Moreover, he
noted that in absence of the index species, the use of
another name for designating the corresponding time
interval is undesirable. In order to support this thesis,
he used the following metaphor (Buckman, 1902,
p. 557): Temp. Edward VII may date an event in
Australia as well as in England, even though Edward
VII never visits Australia.
In his later publications, Buckman (1920, 1927)
began defining infrazonal stratigraphic units (termed
as horizons, faunal horizons, or faunas) based on bra
chiopods. Establishing primarily two such horizons in
STRATIGRAPHY AND GEOLOGICAL CORRELATION
the BathonianCallovian boundary interval of Scot
land (Buckman, 1920), he subsequently increased
their number to eleven based on materials on Dorset
and Yorkshire defining five ammonite horizons in the
same stratigraphic interval (Buckman, 1927).
Unfortunately, Buckman discredited his method
by constructing subsequently incorrect hemerae
sequences based on fossils from other peoples collec
tions and hypothetical interpretations. The reputation
of this method most suffered significantly from criti
cism by Arkell, who was the recognized expert in the
Jurassic ammonite stratigraphy. This researcher
believed that small units such as hemerae cannot be
traced through any spacious regions and are absolutely
inappropriate for correlation, which means that their
defining made no sense (Arkell, 1933, 1956). Time has
proved this outstanding scientist wrong.
As was noted, after Arkells criticism of the Buck
mans method, zonal units were most frequently con
sidered not from positions of maximal resolution in
understanding by W. Waagen, M. Neumayr, and
S. Buckman, but from the standpoint of their correla
tion potential and preservation of the nomenclature
stability (continuity). Being usually potentially divisi
ble, these units represented in fact (and represent until
now) something resembling subsubstages. Mesezhni
kov (1993) termed such an approach as a strati
2
graphic impressionism .
Nevertheless, single or successive infrazonal units
usually termed as horizons gradually received recogni
tion in the Jurassic stratigraphy. This process was
spontaneous, when different criteria were used for
defining new horizons. Of great significance was the
work by Howarth (1958) dedicated to revision of
ammonites from the family Amaltheidae. Although
this author has never defined infrazonal units, he con
sidered in detail peculiar features in evolution of
Amaltheidae representatives paying special attention
to recognition of chronological subspecies, which
characterized successive stratigraphic intervals. Being
a staunch supporter of using the term subspecies
exclusively for geographic varieties, Howarth used in
the situation under consideration the term transient
(this term was proposed by Bather (1927) as an equiv
alent of Waagens mutation). Subsequently, chro
nosubspecies became widely used for defining and
indexing biohorizons.
Callomon (1964, p. 21) was probably the first to
give an extended definition of a horizon as the smallest
infrasubzonal biostratigraphic unit: At the lower
end of divisibility, a stratigraphic unit should meet cer
tain minimum requirements to qualify even as a sub
2 This
(impressionistic) approach itself is inconsistent with
optimization of one of the main stratigraphic tasks such as strati
graphic subdivision and, as follows from the practice, hampers
the solution of another stratigraphic problem, i.e., stratigraphic
parallelization (correlation) decreasing its accuracy due to fre
quently doubtful increase of the correlation potential.
Vol. 20 No. 2 2012
 BIOHORIZONS AS INFRAZONAL BIOSTRATIGRAPHIC UNITS
zone. It should be recognizable over the whole of some
other natural geographical unit, e.g. a whole basin of
deposition, or lithofacies province. As a rough guide
this seems to indicate distances of order of 300 km. A
suitable term for such units is horizon The ultimate
resolvable and observable stratigraphic unit on the
ammonite scale is the thinnest bed or packet of strata
in a single section, which can be identified by a single
species.
During a short period, such horizons were defined
in all the Jurassic stages of different regions. Moreover,
it was considered that the peculiar features of the hori
zons were both their insignificant stratigraphic range
and limited geographic distribution (Callomon, 1994).
As was noted in the review of Jurassic stratigraphy of
France (Mouterde et al., 1971), the zones frequently
correspond to the stratigraphic range of a genus, while
species may be used for more detailed subdivision,
although such units are characterized by the limited
geographic distribution. Such an approach to defining
horizons was also used in the work dedicated to the
Boreal Oxfordian (Sykes and Callomon, 1979).
By the beginning of the 1980s, researchers had col
lected a lot of data on infrazonal successions, which
dictated the necessity in their interpretation. Never
theless, the intense activity of experts in Jurassic
ammonites and stratigraphy was ignored by the Inter
national Commission on Stratigraphy, which practi
cally paid no attention to infrazonal units (see below).
Three works published in the mid1980s played a
key role in development of views on the infrazonal
stratigraphy.
In his article dedicated to problems of bio and
chronostratigraphy, Callomon (1984a) provided a def
inition of a faunal horizon: A faunal horizon is a bed
or series of beds, characterized by a fossil assemblage,
within which no further stratigraphical differentia
tions of the fauna and flora (sic!) can be distin
guished. In the work dealing with the evolution of
ammonites belonging to the family Cardioceratidae
that was published a year later, Callomon (1985, p. 52)
emphasizes the significance of phylogeny as a basis for
infrazonal subdivision, and a horizon is thus defined
is a unit in which no further evolutionary change is dis
cernible and one that may therefore be treated for all
practical purposes as of negligible duration.
When defining faunal horizons, Callomon high
lighted chronological subspecies and suggested using
for their designation nonLinnean symbols such as
3 tons suggested the synonymous term (for a subzone) zonule,
Greek letters . As is shown below, such suggestion
cannot be considered as a fortunate one. As a geochro
nological equivalent of the faunal horizon, Callomon
used the Buckmans term hemera (see above),
which means in Greek a day or time.
3 Greek
letters were used for similar purposes as well (Phelps,
1985).
STRATIGRAPHY AND GEOLOGICAL CORRELATION
215
Phelps (1985) shares different views on the infra
zonal stratigraphy. As a smallest biostratigraphic unit,
this author used a zonule considering it, in fact, as a
subsubzone (in some situations, he defines even
subzonules (Phelps, 1985, fig. 9), although avoiding
4
this term). Phelps noted that the horizon is a very
widely understood term. According to him, the zonule
is a smallest biostratigraphic unit traceable through the
paleobiogeographic province, although its application
may be limited also by a less spacious area with the
high degree of endemism. The zonule should readily
be distinguishable using the stratigraphic range of the
characteristic species. Phelps (1985, p. 343) suggested
a list of subordinate criteria, which may be used for
defining the zonules: (a) in situ transition throughout
the geographical range, i.e. gradualistic chronospeci
ation; (b) quantum evolution of a small population
(with subsequent migration) showing no stratigraphi
cal overlap of parent and daughter species; (c) quan
tum evolution of a small population, with noticeable
overlap of parent and daughter species; (d) acme event
in a shortlived species; (e) acme event in a longrang
ing species. Thus, when defining zonules, the prefer
ence is given to ammonite groups that dwelt in shal
lowwater basins. In general, the stratigraphic resolu
tion is sacrificed to the correlation potential and
zonules became smaller units as compared with zones,
although not necessarily potentially indivisible. Nev
ertheless, when justly criticizing the usage of a hori
zon, which represents as one of the most ambiguous
stratigraphic terms, applied in different meanings, as a
smallest unit, Phelps used a similarly controversial
term. The Fentons (1928) were the first to introduce
the term zonule into the geological practice likely as
a paleoecological, not stratigraphic term. Although
the zonule was accepted as the smallest unit of the
zone first in the North American Code of 1961
5
(Zhamoida et al., 1969) and then in the International
Stratigraphic Guide (1976), its application in the geo
6
logical practice was extremely rare .
The late 20th century was marked by development
of infrazonal ammonite scales practically for the
whole Jurassic System of northwestern Europe and the
Mediterranean region as well as for separate Jurassic
4 Prior to this work by Phelps, the zonules have practically never
been used in the Jurassic stratigraphy. Only Imlay (1984) defined
several zonules in the Bajocian of Alaska, although he likely
considered them as synonyms of subzones.
5 These authors (Zhamoida et al., 1969, p. 56) wrote: The Fen
although the latter is less frequently used. In some works, this
term is used for designating even smaller and more local unit
than the subzone. In England, the similar sense is imparted to
the term horizon.
6 In the sense close to that proposed in the penultimate edition of
the International Stratigraphic Guide (1976), the term zonule
was mainly used by experts in the Quaternary System for desig
nating very small biostratigraphic units. Now, the use of zonules
is admitted to be undesirable (Murphy and Salvador, 1998).
Vol. 20 No. 2 2012
216 ROGOV et al.
intervals of other regions. This resulted in almost an
orderofmagnitude increase of stratigraphic resolu
tion and correlation accuracy, which, in turn, offered
new opportunities for paleontological and geohistori
cal investigations.
Page (1995) summarized the available views on the
usage of the infrazonal method in the Jurassic stratig
raphy. Instead of Callomons term faunal horizon,
7 intervals of geological time. (3) As the boundaries of
this author used in the same sense the term biohorizon .
This author considered the biohorizons and zonules as
principally different infrazonal units (K. Page uses the
term intrasubzonal).
Page (1995, p. 802) considers the zonule as a
smallest component subdivision of a chronostrati
graphical hierarchy, defined, as with higher divisions,
by a basal boundary stratotype, which has nothing to
do with its primary understanding by the Fentons and
is inconsistent with its practical use by Phelps. K. Page
first determined the zonule as a chronostratigraphic
equivalent of the biohorizon together with a potential
hiatus. Later, he turned back to the understanding of
the zonule in (Phelps, 1985). Despite the fact that
K. Page continued considering the zonule as a smallest
unit of the chronostratigraphic hierarchy, in his prac
tical investigations, the latter ceased being the chro
nostratigraphic equivalent of a single biohorizon with
a potential hiatus, but could correspond to several
(sometimes, threefour) successive biohorizons.
Following Callomon, Page (1995, p. 802) deter
mined the biohorizon as a bed or series of beds char
acterized by a fossil assemblage within which no fur
ther stratigraphical differentiation of the fauna (or
flora) can be made, i.e. a biohorizon is effectively
defined at both its base and top in a single reference
section. Such a characteristic makes the biohorizon
different from conventional units of the chronostrati
graphic hierarchy recognizable based on the lower
boundary in a stratotype. The potential discontinuity
of biohorizons is admissible. Such a situation more
adequately characterizes their real range in particular
geological sections, which usually include intervals
that cannot be unambiguously attributed to a certain
biohorizon because of different reasons (rare occur
rence or lack of fossils, their poor preservation, and
others).
Page (1985, pp. 802803) also considered other
important features of biohorizons: (1) The recogni
7 In the International Stratigraphic Guide (1976, 1998) and
Stratigraphic Code of Russia (1992, 2006), the biohorizon is
defined as a certain boundary (datum plane) that has no thick
ness (which is specially emphasized). In such a manner, the bio
horizons are largely used by experts in microfossils. Neverthe
less, the ammonite biohorizons (horizons or faunal horizons)
were long ago introduced into biostratigraphic investigations
mostly of the Jurassic and Cretaceous systems as smallest bios
tratigraphic units, although characterized by some stratigraphic
range in particular sections. In the English guide to stratigraphi
cal procedure (Whittaker et al., 1991), the biohorizon is
accepted precisely in such a sense.
STRATIGRAPHY AND GEOLOGICAL CORRELATION
tion of a biohorizon is intimately related to the identi
fication of the index species/subspecies, the smallest
distinguishable segment of a continuously evolving
lineage. Only the occurrence of diagnostic transient
species can confirm the presence of a particular bioho
rizon, but the general generic or specific composition
of a fauna can be a useful guide to recognition. (2) Bio
horizons represent discrete but typically very short
most successive biohorizons are not coincident, a sig
nificant time gap is potentially present and is shown as
an interval between successive units. (4) A sequence of
biohorizons is established by first constructing a suc
cession of faunal associations, and then distinguishing
geographically persistent or morphologically distinct
associations, (5) The sequence of defined biohorizons
can be integrated with the existing scheme of standard
subzones. Nevertheless, the current state of strati
graphical knowledge inevitably means that boundaries
of biohorizons and subzones may be not coincident.
(6) Biohorizons are usually named in a dual manner,
firstly by consecutive numbering and secondly by
8
selecting a suitable species as an index .
According to Page, correlation between succes
sions of biohorizons provides its higher accuracy as
compared with zonal scale, since (1) Both the bases
and tops of each biohorizon can be correlated, (2) The
intervals between biohorizons can be as important in
correlation as the biohorizons themselves, (3) A bio
horizon can be correlated with a biohorizon plus the
interval below and/or the interval above, or even a
sequence of biohorizons, (4) Following on from the
above, every line on the correlation diagram has a very
specific meaning and may require some discussion in
accompanying text, (5) Exactly correlated biohori
zons form time planes, and can be considered as being
equivalent in event stratigraphy.
Page (1995, p. 805) thoroughly analyzed the nature
of biohorizons defining seven following processes or
factors, which make easier their recognition: (1) The
evolution of one or more ammonite populations/lin
eages within the biogeographical province under study,
leading to the development of distinguishable tran
sients. (2) The evolution and divergence of related
ammonite populations. (3) The evolution of an
ammonite lineage present in the province under study,
or in a separated area of the same province, followed
by the migration of the new form into the study area.
(4) The presence of morphologically complex and
rapidly evolving groups. (5) The migration into the
area being studied, from another basin or province, of
a new ammonite population/lineage. This migra
tion may be shortlived (i.e. one transient/biohorizon
8 The ordered numbers are used for biohorizons only by some,
mainly British researchers. The introduction of ordered indices
(numbers) into names of biohorizons is unreasonable, since it
adds unnecessary detail and the further complication of the
scale.
Vol. 20 No. 2 2012
 BIOHORIZONS AS INFRAZONAL BIOSTRATIGRAPHIC UNITS
duration) or may lead to the establishment of a
local, possibly evolving, population. (6) Changes in
faunal composition including acmeevents (includ
ing dominance) of certain lineages. (7) The periodical
or occasional preservation of suitable ammonite fau
nas resulting from fluctuating sedimentary processes.
According to Page (1995), the duration of zonules
(biohorizons with potential hiatuses) in Jurassic sec
tions ranges from 80 to 722 kyr averaging 200
9 tory of the infrazonal approach in biostratigraphy,
350 kyr . The article by Callomon (1995), which was
published simultaneously with that by Page, is dedi
cated to problems of the infrazonal stratigraphy. Cal
lomon considered any first and last appearance of a
taxon as potential boundaries of biohorizons. Thus,
their ranges (theoretical) were determined by the
composition of guide fossil assemblages, not by com
plete stratigraphic ranges of index species. At the same
time, this author emphasized that such horizons are
not necessarily of practical significance and that of
interest are only horizons, which may be used at least
for correlation within the particular region and, con
sequently, for development of a regional scale.
In his recent work, Meister (2010) demonstrated
the constructive approach to the infrazonal stratigra
phy using the Sinemurian and Pliensbachian stages as
an example. The last researcher considers biohorizons
(for local scales), faunal horizons (for integrated
regional scales), and standard horizons (for scales of
biogeographic provinces and realms) as infrazonal
biostratigraphic units. He understands the zonule as
an infrazonal chronostratigraphic unit distinguishable
by the lower boundary and corresponding to a stan
dard horizon.
Since the 1990s, the infrazonal units became fre
quently used by Russian experts in the Jurassic stratig
raphy (Mitta and Starodubtseva, 1998; Gulyaev and
Kiselev, 1999; Gulyaev, 1999, 2001; Mitta, 1999, 2000;
Kiselev, 2001; Rogov, 2001, and others). In the new
millennium, the development of the infrazonal
approach in biostratigraphy is continued precisely by
Russian researchers. They proposed nomenclature
rules for regulating the recognition, description, and
practical application of biohorizons (Gulyaev, 2002;
Gulyaev et al., 2010) and analyzed their correlation
potential (Rogov et al., 2009). In addition, they con
sidered the biological nature of biohorizons reducing
all their diversity to two main types: phylogenetic and
immigrational (Zakharov et al., 2007).
Similar to the Jurassic System, the infrazonal
approach became widely used in the Cretaceous
(Bulot et al., 1992; Vermeulen, 1997; Hoedemaeker
et al., 2003; and others) and Triassic (Goy, 1995)
stratigraphy. The biohorizons are still distinguished
9 Judging
from recent data, Pliensbachian and Toarcian chrons
may differ in duration from each other approximately by an
order of magnitude (McArthur et al., 2000), while the minimal
duration of hemerae (chronological equivalents of biohorizons)
may be as long as approximately 1020 kyr.
STRATIGRAPHY AND GEOLOGICAL CORRELATION
217
using almost exclusively ammonoids, although there
are examples of their recognition by belemnites
(Mitchell, 2005).
RECOGNITION AND APPLICATION
OF BIOHORIZONS, THEIR CLASSIFICATION,
AND SCALES
The analysis of the more than a century long his
which was particularly consistent and fruitful during
the last three decades, allows the following definition
of a biohorizon to be proposed (we prefer this term as
the most frequently used in the following sense):
The biohorizon is the smallest correlatable biostrati
graphic unit, which encloses a unique integrity of taxa
and is further indivisible based on taxonomic differenti
ation of guide fossils (by phylogenetic and/or immigra
tional events that serve as a basis for biohorizon recogni
tion). The necessary and sufficient condition for recogni
tion of the biohorizon in the particular section is
identification of the species/subspecies with the observ
able stratigraphic range corresponding to this biohori
zon. The remainder of the guide fossil assemblage is of
significance for correlation beyond the distribution area
of the biohorizon.
In other words, the biohorizon is a smallest discrete
correctable unit (sui generis an atom) of biostratigra
phy. Having, by definition, both the lower and upper
boundaries in any section, it principally differs from
conventional units of the stratigraphic hierarchy,
which are and, essentially, go beyond this hierarchy
(for example, one cannot say that zones (subzones) are
divisible into biohorizons or that genera (subgenera)
are divisible into species). From the viewpoint of pre
sumption of isochronisms (Lazarev, 1997), both
boundaries of each biohorizon are principal limits of
accuracy of measurements by the biostratigraphic
method (as minimal divisions of the metrical scale
regardless of its scale). Having a specific status of ele
mentary biostratigraphic units, the biohorizons are
epistemologically defined (separated) according to the
principle of the difference (peculiarity) of biostrati
graphic features; their sequences may, in turn, be
united into larger units by the conventional way based
on the principle of similarity of these features
(Gulyaev, 2002). A century ago, Buckman used a sim
ilar approach in geochronology (see above), when he
defined hemerae subsequently uniting them into ages.
The same approach is used in any hierarchic taxon
omy ensuring its development as a scientific research
program in the terminology by Lakatos (1995).
The main features of the biohorizon are as follows:
(1) indivisibility based on features (phylogenetic or
immigrational events) that serve as a basis for its recog
nition (more detailed, for example, phylogenetic
investigations may provide grounds for its division, not
subdivision of the biohorizon); (2) identification
exclusively based on the index species/subspecies;
Vol. 20 No. 2 2012
218 ROGOV et al.

(3) occurrence of both the lower and upper boundaries
in any section that are exclusively determined by the
observable distribution of the index species/subspecies
(in the optimal case, the boundaries of neighboring
biohorizons in the section coincide between each
other). Inasmuch as these features are separately char
acteristic of other biostratigraphic units, let us pay
attention to their differences from biohorizons.
In some cases, the resolution of current zonal
scales may almost correspond to that of biohorizon
sequences (i.e., zones in them are factually indivisi
ble). At the same time, we should remember that their
construction principles are different. The biohorizons
reflect the biostratigraphic structure of sections more
adequately including also intervals, attribution of
which to any biostratigraphic unit is ambiguous.
Moreover, of correlative significance are both the bio
horizons proper and separating intervals (Page, 1995).
By the principle of their substantiation, the phyloz
ones are most similar to phylogenetic biohorizons. At
the same time, they are not indivisible by definition.
For example, by their ranges the phylozones in the
Jurassic System usually exceed considerably biohori
zons. Such zones are based on the succession of
related ammonite genera or subgenera (for example,
in the Callovian Kosmoceratidae succession), while
biohorizons in this interval are distinguishable based
on successions of species/subspecies.
At the first glance, the biohorizon as a stratigraphic
unit, which may be defined among barren sediments,
slightly resembles the socalled beds with fauna (flora)
(Stratigraficheskii , 2006). Nevertheless, the latter
represent only subsidiary biostratigraphic units and
meet none of requirements to the biohorizons.
As was mentioned, the boundaries of successive
biohorizons may be inconsistent with each other and
potential hiatuses are presented in stratigraphic scales
in form of intervals between neighboring units. This is
natural and convenient, since only some levels sepa
rated by stratigraphic hiatuses or intervals barren of
satisfactorily preserved guide fossils appear to be suffi
ciently characterized by paleontological remains in
most sections (Fig. 3). In this case, it is impossible to
comply with the principle of continuity without spec
ulative assumptions. In some intervals of sections lack
ing visible hiatuses and well characterized by guide
fossils (and well studied) through their entire thick
ness, the observed stratigraphic ranges of successive
species/subspecies belonging to a single phyletic lin
eage may join each other (Figs. 4, 5). In addition, inas
much as the biohorizons are defined using different
principles (evolutionary and migrational) and differ
ent taxa from the guide fossil group, their overlap is
admissible (Figs. 4, 5).
As follows from the practical activity, the biohori
zons are most convenient for correlation within pale
obiogeographic provinces and realms, although some
biohorizons or their sequences may frequently be
traced in several paleobiogeographic realms (Cal
lomon, 2001; Rogov et al., 2009). The correlation
potential of biohorizons is immediately connected
with paleobiogeographic distribution of guide taxa and
their ecological (adaptive) strategy; indirectly, it
depends also on the completeness of the geological
record in different regions.
Page (1995, p. 805) outlined several geological and
biological factors that allow recognition of biohori
zons. The geological factors are primarily related to
depositional environments, postdepositional pro
cesses, and denudation that determine the irregular
preservation and occurrence of fossils. All the diversity
of biological factors outlined by Page may be reduced
to two types: phylogenetic and immigrational. This
provides grounds for defining two corresponding types
of biohorizons (Zakharov et al., 2007):
(1) The phylogenetic biohorizons are distinguishable
by evolutionary events (phyletic speciation) in lineages
of guide taxa that inhabited the region under consider
ation. The basis for defining such biohorizons is
yielded by both stratigraphic investigations proper and
thorough study of the phylogenesis at the species and
subspecies levels. The most frequent result of such
studies is sequences of phylogenetic biohorizons based
on evolutionary successions of guide taxa dominant or
subdominant in the region under consideration
(Fig. 4).
(2) The immigrational biohorizons are recognizable
based on shortterm invasions of certain species/sub
species to the region under consideration (Fig. 5)
(Rogov et al., 2009). Such immigrants usually leave no
descendant, although they may continue evolving in a
new region. Two important features should be noted:
(a) the stratigraphic range of the index species charac
teristic of the immigrational biohorizon may substan
tially be shorter as compared with that of this species
in the eudemic (primary) part of its distribution area;
(b) the same biohorizon may be phylogenetic in some
parts of its area and immigrational in the other one.
In this case, the migration rate may be ignored.
According to the phrase by S. Buckman (19091930,

Fig. 3. The Upper BathonianLower Callovian section on the left side of the Volga River near the Prosek and Isady settlements
of the Lyskovo area in the Nizhni Novgorod region.
Stratigraphic ranges of ammonite assemblages (f1f10), which correspond to biohorizons: (inf) P. infimum, (am) P. cf./aff.
ammon, (jacq) M. jacquoti. (elat) P. elatmase, (sur) C. surensis, (subp) C. subpatruus, (ind) K. indigestus; (curt) L. curtilobus, (gal)
K. galilaeii, (call) S. calloviense (index species in lists are designated by asterisks). The biohorizons are separated either by barren
intervals lacking of reliably identifiable guide forms or stratigraphic hiatuses (wavy lines). The section stratigraphy and identifi
cations of ammonite taxa are given after D.B. Gulyaev. [M] macroconchs; [m] microconchs).

STRATIGRAPHY AND GEOLOGICAL CORRELATION Vol. 20 No. 2 2012

 BIOHORIZONS AS INFRAZONAL BIOSTRATIGRAPHIC UNITS 219

Rondiceras geerzense (Behrendsen) [M] sensu Mitta, 2000

Thickness, m

Gal.
call f10 Novocadoceras sasonovi (Kiselev) [m]

Lithology
Substage

Biohorizon
Pseudocadoceras sp. [m]

Zone
gal f9 *Sigaloceras calloviense (Sowerby) [M]

Bed
Gulielmina quinqueplicata Buckman [m]
Proplanulites (Crassiplanulites?) sp. ind. [M]

Koenigi
curt f8 *Kepplerites (Gowericeras) galilaeii (Oppel) [M]
km2
Rondiceras sokolovi (Kiselev) [M]
K. C.

4 Cadoceras tolype Buckman [M]

ind f7 Cadoceras(?) confusum (Gulyaev) [M]
3
Novocadoceras cf./aff. sasonovi (Kiselev) [m]
subp f6 Pseudocadoceras bellator (Kiselev) [m]
*Kepplerites (Gowericeras) curtilobus (Buckman) [M]
Toricellites curticornutus Buckman [m]
Proplanulites (Proplanulites) ferruginosus Buckman [M]
Subpatruus

P. (P.) cf. excentricus Buckman [m]

sur f5 Chamoussetia chamousseti (dOrbigny) [M]
2g Pseudocadoceras boreale Buckman [m]
Pseudocadoceras aff. grewingki (Pompeckj) [m]
*Kepplerites (Gowericeras) indigestus (Buckman) [M]
Toricellites lahuseni (Parana et Bonarelli) [m]
Proplanulites (Proplanulites) cf. ferruginosus Buckman [M]
P. (P.) cf. excentricus Buckman [m]
*Cadochamoussetia subpatruus (Nikitin) [M]
Kepplerites (Gowericeras) cf. unzhae Gulyaev [M]
Lower Callovian

2f Homoeoplanulites sp. ind. [m]

*Cadochamoussetia surensis (Nikitin) [M]
elat f4 Cadoceras sp. ind. [M]
Pseudocadoceras aff. mundum (Sasonov) [m]
Kepplerites (Gowericeras) unzhae Gulyaev [M]
Toricellites pezhengensis Gulyaev [m]
Macrocephalites (Macrocephalites) pavlowi Smorodina [M]
Elatmae

2e Homoeoplanulites spp. [M, m]

*Paracadoceras (Rossicadoceras) elatmae (Nikitin) [M]
Cadoceras quensledti simulans Spath [M]
Pseudocadoceras mundum (Sasonov) [m]
Macrocephalites (Macrocephalites) prosekensis Gulyaev [M]
M. (M.) volgensis Gulyaev [M]
2d M. (M.) verus Buckman [M]
2c M. (Pleurocephalites) cf. terebratus (Phillips) [M]
jacq 2b f3 M. (Kamptokephalites?) cf. zickendrathi Mitta [m]
2 M. (K.?) sp. juv. [m?]
am f2 Paracadoceras (Rossicadoceras) ex gr. poultoni Gulyaev [M]
Pseudocadoceras sp. nov. (aff. pisciculus (Gulyaev)) [m]
1f Kepplerites (Kepplerites) sp. ind. [M]
Toricellites sp. ind. [m]
*Macrocephalites (Macrocephalites) jacquoti (H. Douville) [M]
inf 1d f1 *Paracadoceras (Catacadoceras) cf./aff. ammon (Spath) [M]
Pseudocadoceras pisciculus (Gulyaev) [m]
Upper Bathonian
Infimum

Kepplerites (Kepplerites) sp. ind. [M]

1c Toricellites cf./aff pauper (Spath) [m]
*Paracadoceras (Catacadoceras) infimum (Gulyaev et Kiselev) [M]
Pseudocadoceras pisciculus (Gulyaev) [m]
Kepplerites (Kepplerites) svalbardensis Sokolov et Bodylev [M]
1b Toricellites pauper (Spath) [m]
scale 1 m

1
Vertical

Tatarian
Stage

STRATIGRAPHY AND GEOLOGICAL CORRELATION Vol. 20 No. 2 2012

220 ROGOV et al.

Thickness, m
Substages, Lithology Phyletic lineage
Biohorizons of the genus Funiferites
zones, subzones
Oxfordian

Cor Cordatum
datum Costicardia
Lower

Mariae
Lamberti

1 cm
Spinosum K. kuklikum
Funiferites funiferus

s
ru
ife
fun
F.
K. rowlstonense

F. patruus

Proniae
Upper Bathonian

Athleta

K. proniae

1 cm
F. sp. nov.

Funiferites patruus
K. phaeinum

F. allae

Phaeinum

1 cm
Funiferites sp. nov.
oronatum

K. grossouvrei
Middle Callovian

Grossouvrei
K. posterior
Vertical scale 1 m

Obductum K. crassum
K. jason jason
Jason
1 cm
Jason

Medea K. medea medea Funiferites allae

Fig. 4. The MiddleUpper Callovian to Lower Oxfordian section in the Mikhailovtsement quarry near Mikhailov of the Ryazan
Oblast, after (Kiselev et al., 2003) modified.
Dark gray coloration indicates stratigraphic ranges of evolutionary successive species from the ammonite genus Funiferites (Car
doceratidae) and corresponding biohorizons; light gray coloration shows the standard sequence of biohorizons based on the
stratigraphic distribution in the phyletic lineage of the ammonite genus Kosmoceras (Kosmoceratidae).

pt. XXXVII, p. 24), the rate of ammonite migration units. In the practical activity, such a situation is rare
to that of deposition was like the flight of an aeroplane because of insufficient knowledge or incomplete geo
to the progress of bricklaying. logical record. The complete sequence of biohorizons
In the optimal case, the sequence of biohorizons in a single section cannot usually be established for dif
exactly corresponds to the previously established zonal ferent reasons and for constructing the continuous

STRATIGRAPHY AND GEOLOGICAL CORRELATION Vol. 20 No. 2 2012

 BIOHORIZONS AS INFRAZONAL BIOSTRATIGRAPHIC UNITS 221

Ammonite ranges Appearance/

Paralingulaticeras (Rogoviceras) cf. efimovi

Thickness, m
disappearance levels of

Subzone

Lithology
Substage
Zone Biohorizon taxa (potential

Paralingulaticeras (Rogoviceras) efimovi

boundaries of
assemblagebased

Bed
stratigraphic units)
Tenuicostatum
Klimovi Sokolovi Pseudoscythica

1/67

Franconites spp.
P. puschi

Ilowaiskya pavida

Sphinctoceras spp.
1/8a, b

Ilowaiskya sokolovi

?Fontannesiella sp.
?Danubisphinctes sp.
Ilowaiskya klimovi
S. neoburgense

Eosphinctoceras sp.
Pseudovirgatites puschi
1/9ac

Sutneria asema
Lower Volgian

I. pseudoscythica

Pseudovirgatites tenuicostatum

Schaireria neoburgense
1/10
P.

Franconites 1/11

Aulacostephanus spp.
I. pavida 1/12

Ilowaiskya pseudoscythica
I. sokolovi
P. efimovi 1/13a, b
I. klimovi

N. steraspis 1/14

N. cf. praecursor 1/15

1/1618

Neochetoceras steraspis
Neochetoceras cf. praecursor
943=

Taramelliceras cf. franciscanum

Ochetoceras cf. zio
S. ilowaiskii 1/19
Fallax

9/3942
Sarmatisphinctes ilowaiskii

9/38
S. fallax

Neochetoceras rebouletianum
Sarmatisphinctes subborealis

Suboxydiscites cf. taimyrensis

9/3637
9/3435
Sarmatisphinctes fallax

9/3233
9/2931
S. zeissi
9/28
Autissiodorensis
Upper Kimmeridgian

9/2527
Sarmatisphinctes cf. subborealis

Nannocardioceras sp.
Sarmatisphinctes zeissi

9/2324
Subborealis

Neochetoceras cf. subsidens

9/22
Nannocardioceras volgae
Nannocardioceras krausei
Nannocardioceras cf. anglicum

S. subborealis 9/1921
Sutneria aff. rebholzi

9/1418
Schaireria sp.

9/13
9/12
Neochetoceras ex gr.

N. volgae 9/1011
S. aff. rebholzi 9/9
Contejeani

scale 1 m

subnudatum
Eudoxus

9/46
Vertical

N.anglicum
9/3

Fig. 5. The Upper KimmeridgianLower Volgian section near the Gorodishchi village in the Ulyanovsk raion of the Ulyanovsk
Oblast.
The distribution of ammonites and infrazonal subdivision of the section is shown after (Rogov, 2010) modified. The stratigraphic
ranges of ammonite, the phyletic lineages of which are used for constructing the main scale of biohorizons (Sarmatisphinctes
IlowaiskyaPseudovirgatites) are shown in black; stratigraphic ranges of other ammonite species are given in gray. The phylo
genetic and immigrational biohorizons are shown in white and gray, respectively. (P.) Pseudoscythica.

STRATIGRAPHY AND GEOLOGICAL CORRELATION Vol. 20 No. 2 2012

 222

Regional Stratigraphic structure of local sections (see numerals in the map)

scale (1) Churkinskaya
(2) Pezhenga
(3) Prosek
(4) Khvadukasy
( (5) Lazarevka
(6) Uzhovka
(7) Malinovyi
Shchelya Isady Yazykovo Ovrag Naryan
R.

Biohorizons Mar

Substage
Biohorizons (gray), beds with fauna, gaps in observations, hiatuses

Zone
Ammonites are
Hiatus
Pechora

Ch. crobyloides are not found *Ch. crobyloides

Hiatus Hiatus Hiatus M
ez
en
C. uzhovkensis *C. uzhovkensis R? 1
Hiatus ?
Hiatus Hiatus
C. subpatruus C. subpatruus *C. subpatruus
Beds with Beds with
Cadochamoussetia Cadochamoussetia
and Kepplerites unzhae

Subpatruus
C. surensis C. surensis C. surensis *C. surensis
Se

.
ve

aR
rn

in
ay

C. tschernyschewi C. tschernyschewi *C. tschernyschewi Hiatus Hiatus Hiatus ? Hiatus Hiatus ?

a

Dv
Syktyvkar
P. elatmae P. elatmae P. elatmae *P. elatmae P. elatmae Ammonites are P. elatmae P. elatmae
not identified
Ammonites are Ammonites are

Lower Callovian
P. chvadukasyense *P. chvadukasyense not identified P. chvadukasyense P. chvadukasyense not identified
Inaccessible Ammonites are Ammonites are
for observation not identified not identified 2
P. primaevum *P. primaevum Ammonites are

Elatmae
Ammonites are not identified
Hiatus

M.
M.
not identified
P. poultoni *P. poultoni

jacquoti
jacquoti
M. jacquoti M. jacquoti M. jacquoti
P. cf./aff. P. cf./aff. ammon P. cf./aff. ammon P. cf./aff. ammon P. cf./aff. ammon
ammon Coastalmarine Vol
ga Kazan
ROGOV et al.

(subcontinental) R
Inaccessible 3 4
P. infimum P. infimum *P. infimum for observation P. infimum facies Hiatus Nizhni R.

Infimum
a
Novgorod 5 Kam
Inaccessible Beds with Inaccessible 6
for observation Kepplerites (Kepplerites) for observation Coastalmarine
and Cadoceratinae (subcontinental) Samara
facies
Hiatus

Upper Bathonian
R.
lga
Vo N
7
Samara

STRATIGRAPHY AND GEOLOGICAL CORRELATION

S
Do
nR
.
150 km
Volgograd

Vol. 20
Fig. 6. The regional scale of biohorizons in the BathonianCallovian boundary interval of European Russia based on correlated sequences of biohorizons in local sections (after

No. 2
Gulyaev and Kiselev, 1999; Gulyaev, 1999, 2001, 2005, 2007, 2011; Gulyaev et al., 2002). The scale is based on the phyletic lineage of endemic Paracadoceras (P.)Cadoch
amoussetia (C.)Chmoussetia (Ch.) representatives; in addition, the scale includes the biohorizon defined by the Tethyan immigrant Macrocephalites (M.) jacquoti and repre
senting an interregional biostratigraphic reference unit. The biohorizons are shown in gray. Asterisks designate stratotypes of biohorizons.

2012
 BIOHORIZONS AS INFRAZONAL BIOSTRATIGRAPHIC UNITS
regional scale of biohorizons, one should correlate
their successions in a series of sections scattered some
times through a relatively spacious region (Fig. 6). The
thorough reconstruction of phyletic lineages among
guide fossils may be helpful in such correlations.
When zonal subdivision and recognition of bioho
rizons is based on different guide fossil groups, the bio
horizon may overlap boundaries of zonal units. In the
same stratigraphic interval of a single region (deposi
tional basin, paleobiochorema), parallel sequences of
biohorizons may be established using different phyl
etic lineages of guide fossils or invasions of migrating
species. The distribution areas of such biohorizons and
their sequences are usually not coincident. At the
same time, in areas of their cooccurrence, these
sequences may be integrated into a single complex
scale, which offers wide opportunities for interre
gional correlations. It is understandable that bound
aries of parallel biohorizons in the integrated scale may
not necessarily be coincident (Fig. 7).
NOMENCLATURE AND DESCRIPTION
OF BIOHORIZONS
The rapid development and frequently spontane
ous application of the infrazonal method against the
background of its relatively poor theoretical substanti
ation resulted in the unavoidable anarchy in defining
biohorizons.
The thoughtless approach to index taxa became
one of main negative trends. This trend was initiated
by Callomon (1985), who suggested designating the
chronological index species of consecutive biohori
zons lacking proper names by nonLinnean sym
bols, i.e., by Greek letters (for example, Cadoceras
apertum , , ). Subsequently, some researchers used
Roman or Arabic numerals for the same purpose
(Dietl, 1994; Page, 2003; and others). In addition, the
species in the open nomenclature or only genera
(sometimes, with additional numerals or letters) are
frequently used for indexing the biohorizons. It is clear
that such an approach introduces the ambiguity in the
understanding of the defined stratigraphic units,
which should be considered only as conditional (pre
liminary) units that need to be designated by a suitable
index species/subspecies.
In some works (Scweigert, 1998; Page, 2003; Meis
ter et al., 2005; and others), two (less commonly,
three) species from the guide fossil group are used for
indexing the biohorizons. In such situations, one (two)
index species appears usually to be superfluous, since
its stratigraphic range coincides with that of the other
index species or exceeds the latter. The recognition of
biohorizons based on the partial overlap of strati
graphic ranges of several species is unreasonable, since
such an approach may potentially result in recognition
of many unimportant stratigraphic units with the dis
tribution area in common and practically without any
correlation potential (see appearance/disappearance
STRATIGRAPHY AND GEOLOGICAL CORRELATION
223
levels of taxa in Fig. 5). In such a situation, most suit
able are parallel sequences and integrated scales of
biohorizons (see above).
In some cases, the biohorizons are distinguished by
the acme level of the index species (Page, 1995). Such
an approach should also be viewed as undesirable
because of the doubtful correlation potential of corre
sponding stratigraphic units, since (1) no uniform cri
teria exist for boundaries of the acme level and
(2) acme levels of the same species in different areas
even of the same basin may be not coincident (for
example, due to ecological reasons).
The recognition of biohorizons without the refer
ence to the particular section interval or, even, the
whole section, where they are defined, became the
second undesirable trend in stratigraphy (Mitta and
Starodubtseva, 1998; Kiselev, 2005; and others). When
the researcher has no chance to illustrate in his work
particular sections and stratigraphic intervals or refer
to their description in earlier publications, he should
operate with faunal assemblages (faunas) similarly to
Callomon (1984a, 1993).
Some researchers define series of horizons in
irregularly condensed sediments by grouping the co
occurring, although presumably differentage species
into consecutive assemblages. Moreover, the fact that
the term horizon implies a certain stratigraphic
range is absolutely ignored. In such a situation, we
should speak about provisional faunal assemblages.
The existing nomenclatural ambiguity determined
the necessity in development of a code of basic rules
for recognition and description of biohorizons
(Gulyaev, 2002; Gulyaev et al., 2010). In absence of
such a code, further investigations can discredit the
infrazonal method. In this work, we propose a prelim
inary version of such nomenclatural rules based on
materials that were accumulated after works by Cal
lomon (1984a, 1985), which triggered rapid develop
ment of the infrazonal ammonitebased biostratigra
phy.
The availability and validity are most important
characteristics of any taxon, the stratigraphic unit
included.
The availability is determined by several criteria.
(1) The name of a biohorizon should include sev
eral components: (1) the word biohorizon or its
replacing synonym (faunal horizon, horizon); the use
of its index number in the regional/local succession is
unreasonable; (2) the name of a single suitable index
taxon (species or subspecies) in its short or complete
form without indication of the author; the use of non
Linnean symbols (letters, numerals) as well as selec
tion of species in the open nomenclature, genera, or
yet not described taxa as indices are inadmissible: all
such characteristics should be considered as condi
tional (preliminary) and require further specification;
(3) the Latin name of the author and year of the bio
horizon name publication (establishment). The last
Vol. 20 No. 2 2012
 Integrated regional
224

Subzone scale of biohorizons C a r d i o c e r a t i d a e

Zone

Stage
Oppeliidae
Cordatum C. cordatum

Subzone
Brief mass immigration of C. (C.) cordatum
Costicardia C. costicardia Subtethyan taxa C. (C.) costicardia

Bukowskii T. baccatum Taramelliceras baccatum C. (S.) bukowskii

C. bukowskii

Cordatum
C. praecordatum C. (S.) praecordatum

Lower
Praecordatum C. alphacordatum C. (S.) alphacordatum

Oxfordian
C. (S.) sp.
P. praemattini [m=Protocard. praemartini]

Mariae
C a r d i o c e r a s
Scarburgense C. scarburgense Kosmoceratidae C. (S.) scarburgense
K. (M.) zudacharicum Q. pseudolamberti
Q. paucicostatum [m=Q. paucicostatum]
K. mojarovskii K. (M.) mojarovskii ?
Lamberti Q. lamberti Q. lamberti

Lamberti
Q. praelamberti K. (L.) geminatim Q. praelamberti

a
Henrici Q. henrici Q. henrici

Quenstedtoceras (Scarburgiceras) (Cardioceras)

Funiferites

Upper
r
Spinosum K. kuklikum K. (L.) kuklikum ?
K. rowlstonense F. funiferus K. (L.) rowlstonense L. keyserlingi F. funiferus

e
Proniae F. patruus F. patruus
K. proniae F. sp. nov. K. (L.) proniae ?

Athleta
F. sp. nov.
Phaeinum K. phaeinum K. (Z.) phaeinum L. lahuseni
F. allae F. allae

o
K. grossouvrei K. (Z.) grossouvrei ?
Grossouvrei ?

m
L. alpha
ROGOV et al.

K. posterior K. (Z.) posterior

s
K. crassum K. (Z.) crassum
L. stenolobum L. stenolobum P e r i s p h i n c t i d a e
Obductum

Coronatum
Longaeviceras
K. obductum K. (Z.) obductum

o
L. praestenolobum L. praestenolobum

Callovian
K. jason jason K. (G.) jason jason R. ? sp. nov.

iddle
K
Jason Pseudoperisphinctinae
K. jason
sedgwickii K. jason sedgwickii

Jason
K. medea magnum K. (G.) medea magnum
Medea R. milaschevici R. milaschevici
milaschevici K. (G.) medea medea milaschevici
K. medea medea

(Gulielmites) (Zugokosmokeras)(Lobokosmokeras) (Mojarovskia)

Anaplanulites submutatus

STRATIGRAPHY AND GEOLOGICAL CORRELATION

C. enodatum aeeta C. enodatum aeeta
C. enodatum R. milaschevici R. milaschevici
enodatum khudyaevi C. enodatum planicerclus khudyaevi
Enodatum Proplanulitinae
C. enodatum Anaplanulites difficilis

ni
C. enodatum enodatum

Cata
planicerclus A. difficilis

ki

Vol. 20
R. tcheffkini

eff .
sigaloceras

h
P. cracoviensis C. pagei Poplanulites cracoviensis

tc R
C. pagei Anaplanulites sp.
R o n d i c e r a s

Calloviense
S. micans S. micans

Lower (pars)
Calloviense R. geerzense
Poplanulites ex gr. petrosus

No. 2
S. calloviense S. calloviense (sensu Mitta, 2000)
ceras
Sigalo

2012
 BIOHORIZONS AS INFRAZONAL BIOSTRATIGRAPHIC UNITS 225

Fig. 7. The integrated scale of Callovian and Lower Oxfordian biohorizons for European Russia based on parallel phyletic lineages
of lowboreal kosmoceratids (gray) and highboreal cardioceratids (dark gray), the geographic distribution areas of which signif
icantly overlap each other (after Kiselev, 2001, 2005, 2006; Gulyaev et al., 2002; Kiselev et al., 2003; Kiselev and Meledina, 2004;
Kiselev and Rogov, 2005; Gulyaev, 2005; Kiselev et al., 2006; Wierzbowski and Rogov, 2011).
The scale includes the immigrational biohorizon based on the subTethyan Oppelidae representative, which is of particular
importance for correlation. The biohorizons based on subboreal Perisphinctidae species are largely used in areas, where kos
moceratids are rare or not found. Horizontal and vertical arrows show immigrations and phylogenetic connections of ammonites,
respectively. The dark gray and light colors designate biohorizons based on highboreal cardioceratids and lowboreal kosmocer
atids, respectively. Other biohorizons are given in white.

component is indicated in revisions and descriptions.
The author of the biohorizon is the researcher, who
first established the corresponding stratigraphic unit
precisely as biohorizon, even though its index coin
cides with that of previously defined host zone or sub
10
zone. The year of publication of the abovemen
tioned work by Buckman (1893) is considered as a
starting date for availability of biohorizon names.
(2) A new name of the biohorizon as well as any
nomenclatural operation or information that may
influence the nomenclature should be published in line
with same rules, which are accepted in current inter
national codes of the zoological and botanical nomen
clatures,
(3) An index of the biohorizon should have a suit
able, although no necessarily valid name. To avoid dis
crepancy, the first description of the biohorizon should
be accompanied by the reference to the illustration
and description of the index taxon as well as by illus
tration of available specimens, which presumably
belong to the latter.
(4) Any biohorizon should have a stratotype. Its
main purposes are (a) to protect the nomenclatural
name (stability of the nomenclature) and (b) to pro
vide potential falsifiability (testing) of initial data. In
addition to the stratotype, the description of the bio
horizon should be provided with indication of the stra
totype area, where this unit bears similar paleontolog
ical and geological characteristics. In case of the loss of
a holostratotype or impossibility of its establishment,
the latter may be replaced with a lectostratotype.
Beyond the stratotype area, stratigraphic standards
(reference sections) may be selected. The latter bear
no nomenclatural significance and provide an addi
tional characteristic of the biohorizon in the study
region. It is desirable that the stratotype section in the
stratotype area represented also the stratigraphic stan
dard. When the lectostratotype cannot be defined in
the stratotype area, any of the stratigraphic standards
may be defined, if needed, as a neostratotype.
Validity. In the zoological and botanical nomencla
tures, the principle of priority, application of which is
reduced only to the seniority of the suitable name pub
10In some works (Hantzpergue, 1989; Biostratigraphie , 1997),
the authors of senior zonal units are considered, through a mis
understanding, as authors of synonymous biohorizons; such an
approach is unacceptable because of principal differences
between these categories of stratigraphic units.
lication, serves as a main criterion of the validity. In
case of biohorizons, it is reasonable to use the triple
priority principle that includes subordinate principles
of (1) minuteness, (2) continuity, and (3) priority.
(1) The principle of minuteness is most important
among characteristics following from the biohorizon
definition proper. According to this principle, the bio
horizon with the shortest geochronological range is a
senior unit relative to its counterparts with wider geo
chronological ranges (for example, three biohorizons
instead two in the same interval).
(3) The principle of continuity means that the bio
horizon defined by the species (subspecies) of the
same phyletic lineage characteristic of their neighbor
ing counterparts is senior relative to the biohorizon
defined by the species (subspecies) from the other
phyletic lineage, if this is consistent with the minute
ness principle. It is natural that evolutionary phases
(cycles) in different phyletic lineages may be not coin
cident; in such cases, the purpose of the continuity
principle is to avoid the potential overlap of strati
graphic units. If neighboring biohorizons are defined
by species from different phyletic lineages, but their
ranges are similar, the name of the middle biohorizon
should be selected by the conventional way, although
there are always some preferences. In case of the avail
ability of parallel phylogenetic and immigrational bio
horizons, it is reasonable to keep both types of strati
graphic units within a single scale, since phylogenetic
biohorizons reflect in the most degree the continuity
principle, while immigrational biohorizons are usually
characterized by the higher correlation potential.
The principle of priority means that the oldest suit
able name of the biohorizon is senior relative to its
later names, when this is not inconsistent with the
principles of minuteness and continuity.
It should be kept in mind that the nomenclature of
biohorizons is to a certain extent dependent on the
nomenclature of index species.
(1) When the name of the index species is changed
because of objective reasons (objective synonymy or
homonymy), i.e., the nomenclatural type is preserved,
the name of the biohorizon becomes also automati
cally changed, although the author remains the same
and its last name is taken into parentheses.
(2) When the name of the index species becomes
invalid due to subjective synonymy, the name of the
biohorizon becomes invalid as well. This is explained

STRATIGRAPHY AND GEOLOGICAL CORRELATION Vol. 20 No. 2 2012

226 ROGOV et al.
by the fact that stratigraphic ranges of the senior and
junior synonyms may potentially be different. The
authorship of a new name should belong to the
researcher, who was the first to revise the latter. If the
geochronological range of the biohorizon and its posi
tion in the scale remain unchanged, the initial name is
included into the synonymy of a new name.
Following below is the recommended plan for
description of biohorizons (n/o means a not obliga
tory item).
Name (with the authors name, year of publica
tions, and a mark nov. for new biohorizons).
Synonymy (with indication of reasons responsible
for their rejection for invalid names).
Index (with the reference to the illustration and
description).
(Stratotype (with the obligatory indication of a par
ticular interval in the published section).
Stratotype area (n/o).
Stratigraphic standards (n/o).
Paleontological characteristic (position of the bio
horizon in the local scale and correlation with scales
available for other regions).
Remarks (n/o).
Distribution (regions, where this horizon is distin
guishable).
Locality (the list of sections, where this biohorizon
is recorded by the author).
We are far from the thought that the above list of
nomenclature rules takes into consideration all the
possible situations and eliminates all the inconsisten
cies. In fact, we had no such a purpose. In our opinion,
it is of importance that this list may serve as a basis for
further developments, investigations, and discussions
among interested researchers. The proposed princi
ples of recognition and application of biohorizons
were recently successfully used in detailed investiga
tions of BathonianCallovian (Gulyaev, 1999, 2001,
2005; Kiselev, 2001, 2006; Kiselev and Rogov, 2007;
and others) and KimmeridgianVolgian (Rogov,
2005, 2010) sections.
CONCLUSIONS
The use of biohorizons opens new prospects for
solving two main stratigraphic problems: subdivision
and correlation. Owing to infrazonal units, the solu
tion of both these problems becomes progressively
more easier. It is not incidental that most discussions
dedicated to the GSSPs of Jurassic stages are reduced
to the question: the base of what biohorizon should be
used for placing a stratigraphic boundary.
It seems that world infrazonal stratigraphy is now at
the development stage even for the Jurassic System.
We hope that the ideas discussed in this work may help
researchers, who are engaged in solving stratigraphic
problems of the Jurassic and other systems based on
ammonites and other fossil groups, in more detailed
STRATIGRAPHY AND GEOLOGICAL CORRELATION
subdivision and correlation of Phanerozoic succes
sions using the experience available from the study of
the Jurassic System. Such examples are known: the
biohorizons are used with progressively growing fre
quency in investigations of the Cretaceous and Triassic
systems; belemnitebased infrazonal scales are also
becoming available. Thus, we may expect the further
progress in development of detailed biostratigraphic
investigations.
ACKNOWLEDGMENTS
We are grateful to our colleagues I.S. Barskov,
V.N. Beniamovsky, A.Yu. Gladenkov, Yu.B. Gladen
kov, V.A. Zakharov, O.A. Korchagin, V.V. Mitta,
S.V. Naugolnykh, V.A. Prozorovskii, D. Bert,
M. Hart, J.L. Latil, F. Olriz, and K. Page for joint
discussions of the main concepts of this work as well as
to its reviewers B.N. Shurygin and E.Yu. Baraboshkin
for their valuable recommendations. This work was
supported by the Russian Foundation for Basic
Research, project no. 090500456.
Reviewers E.Yu. Baraboshkin and B.N. Shurygin
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