Proterozoic Stromatolite Taxonomy and Biostratigraphy

Mikail A. Semikhatov, Maria E. Raaben

Geological Institute, Russian Academy of Sciences, Pyzhevsky 7, Moscow 109017, Russia

Abstract. Precambrian stromatolites are unique objects in Earth mental setting. At the same time, impressive results in
history. The predominance of microbes in ecosystems that they the study of Holocene, predominantly intertidal, stro-
document, their specific global environments, and the scale of their matolites (e.g. Logan et al. 1964; Walter 1976) brought a
evolution have no counterparts in the Phanerozoic. Among several,
basically different stromatolite classifications known in the litera-
group of influential workers to the conclusion that all the
ture, the current version of the traditional system is most exten- above problems regarding Precambrian forms can and
sively employed in the study of Precambrian buildups.lt is artificial should be resolved by means of comparative actualistic
in nature but follows conventional rules of paleontological classifi- analysis. As a result, two schools of stromatolite workers
cation and requires definition of a hierarchy of taxa: forms (form- -biostratigraphic and environmental ones - arose in the
species), groups (form-genera), and types. At present, there is a
1960s. Mechanisms of stromatolite formation, the range
common understanding of stromatolite characteristics, providing
the most efficient basis for definition and identification of the tra-
of settings inhabited by Proterozoic forms, the degree of
ditional system taxa. The types are based on the most general fea- environmental impact on morphology, as well as the
tures of the buildup's morphology. Groups are based on particular very necessity and required resolution of stromatolite
combinations of morphological characteristics defined by the classification were all treated differently by the two
mode of accretion and shape of stromatolite laminae (plus some schools in the 196os and 1970s. Subsequently, contradic-
general features of the microstructure in several cases). Forms are
tions were partly smoothed-out, but progress in under-
predominantly or exclusively based on microstructure. The strati-
graphic potential of Precambrian stromatolites, revealed by empir- standing the morphogenesis, environmental setting and
ical time-and-space distribution data of the distinctive assem- stratigraphic potential of Proterozoic stromatolites was
blages, is evident. Stromatolites are not suitable for the subdivision ultimately based on empirical data concerning their dis-
of the Proterozoic, but provide paleontological characterization of tribution in long-ranging successions, whereas the di-
units which have been defined by other methods and significantly rect actualistic interpretation of Precambrian forms
contribute to their correlation especially within the limits of partic-
ular stromatolite provinces. Interprovincial stromatolite-based
gave rise to several, at one time widely accepted, but mis-
correlations are of lower reliability and time-resolution due to leading dogmas (see Serebryakov 1975; Monty 1977 for
strong variations in the taxonomic composition of coeval stromat- discussion).
olite assemblages across provincial boundaries. Precambrian stro- Pioneer attempts to understand the stratigraphic
matolites demonstrate distinctive directional secular changes in potential of Precambrian stromatolites were made by
taxonomic composition and diversity which were defined by the Walcott, the Fentons, Maslov and others as far back as
evolution of both global environmental and biological factors rele-
vant to the construction and habitat restrictions of these biolites. the 1910s-1940s (see Maslov 1960, for review), but it was
not until the 1960s-1970s that the first straightforward
relevant results were obtained by a number of Russian,
1 Australian, American and French workers (see Raaben
Introduction 1969; Semikhatov 1976, 1991; Preiss 1977; Bertrand-Sar-
fati and Walter 1981 for review). Progress in this field
The Proterozoic was the "golden age" of stromatolites. was related to the development of an improved classifi-
During this extended period of time they reached a max- cation and taxonomy of Precambrian stromatolites and
imum in abundance, diversity, and lateral environmen- the recognition of directional secular changes among
tal expansion. By the very end of the Precambrian, stro- their lower taxa, firstly in northern Eurasia and, later
matolites had declined in response to important ecosys- on, elsewhere.
tem changes (Walter and Heys 1985; Awramik 1991; Grot-
zinger 1990, 1994; Sochava and Podkovyrov 1992; Semik- 2
hatov and Raaben 1994, 1996; Fedonkin 1996, and refer- Taxonomy
ences therein). The stromatolite buildups were (and are)
often regarded as representing a self-sufficient model for Disagreements on principles of classification and tax-
resolving general problems of their morphogenesis, onomy have always been an impediment to the under-
classification, stratigraphic significance and environ- standing of the biostratigraphic potential of Protero-

R.E. Riding and S.M. Awramik (Eds.)

Microbial Sediments
© Springer-Verlag Berlin Heidelberg 2000
296 M.A. Semikhatov, M.E. Raaben

zoic stromatolites. The very nature of stromatolitic 2.2

constructions even gave rise to the opinion that their Form-Genera and Form-Species
morphology was under complete environmental con-
trol and that they did not need to be classified. How- In the infancy of the system, when genera were consid-
ever, most workers admitted the necessity of stromato- ered to be the main stromatolite taxa, some authors dis-
lite classification for the sake of rapid and adequate tinguished them according to both the "mode of
identification leading to their employment as biostrati- growth" of the laminae and the configuration of the
graphic or environmental indicators. It is evident that structures (e.g. Pia 1927), whereas others concentrated
biostratigraphic and environmental disciplines require on the lamina shape alone (e.g. Fenton and Fenton 1937;
classifications of differing resolution. Nevertheless, it Maslov 1939). Such a concentration of emphasis re-
was generally accepted that classifications are artificial sulted in a swelling of the genus Collenia, so that it en-
and should be based primarily upon gross morphol- gulfed almost all other genera. More recently, the em-
ogy. The largest taxonomic categories of stromatolites, ployment of a number of stromatolite characteristics
however one names them (e.g. Korolyuk 1960; Krylov has led to a more restricted notion of what constitutes
1963, 1975; Logan eta!. 1964), are based on the principal a genus.
characteristics of morphology. This level of classifica- Bertrand-Sarfati and Walter (1981) considered the
tion is adequate for the purposes of environmental in- current state of stromatolite systematics to be confused
terpretation. However, Precambrian biostratigraphy and referred to Krylov's (1975, 1976) opinion that there
requires a classification of higher resolution. were 12 separate classifications exploring different di-
agnostic features and overlapping to varying extents.
2.1 However, these classifications (or more precisely, as
Traditional System recognized by Krylov, different approaches to stromat-
olite classification) are by no means competing, con-
The literature contains descriptions of about 1500 stro- currently used systems. The differences between them
matolite taxa of various ranks. About 90% of them have are not related to dissimilar principles of classification,
been designated within the framework of the tradi- but mainly concern either integral properties of build-
tional system. This is artificial in nature, but follows ups of different types (e.g. stratiform and columnar),
conventional rules of paleontological classification: transfer from an original to an amended definition of a
recognition of a number of hierarchic taxa, usage of bi- taxon, or, to a lesser extent, dissimilar estimations of
nomial Linnean nomenclature, formal definition of the taxonomic importance of a diagnostic feature.
type form and type species (holotype), and acceptance Although several workers have advocated the use of
of the priority principle. any outstanding morphological and/or microstruc-
Such an approach originated when stromatolites tural characteristic of buildups for the definition of
were considered as blue-green algae or primitive ani- stromatolite genera (e.g. Walter 1972; Bertrand-Sarfati
mals. The first step in elaborating a separate stromato- 1972; Preiss 1972; Bertrand-Sarfati and Walter 1981;
lite system was made by Pia (1927), who subordinated Grey 1984), most of the valid taxa of generic rank are
previously described genera to the new higher-rank based on morphological characteristics (see Krylov
taxon Stromatolithi Pia and treated these genera as 1963, 1975, 1976; Raaben 1969; Semikhatov 1976, 1991,
conventional taxa, representing growth-types of a sub- for review) that are defined by the mode of accretion
family of blue-green algae. Understanding of the real and shape of stromatolitic laminae (Hofmann 1969;
nature of stromatolites gave rise to debate concerning Raaben 1969). A suggestion to consider peculiarities of
the appropriateness and rationale of applying biologi- microstructure as an obligatory attribute of form-gen-
cal nomenclature to what are in fact organo-sedimen- era (e.g. Komar 1966) has not gained widespread accep-
tary structures (e.g. Cloud 1942; Maslov 1960; Hofmann tance.
1969). As a result, several stromatolite systems have At present, there is a fairly uniform understanding
been suggested as alternatives to the traditional one of characteristics providing the most useful basis for
(see Hofmann 1969; Walter 1972, for review), but an im- definition and identification of stromatolite form-gen-
proved version of the latter, based on newly recognized era. For stratiform and nodular buildups these are the
morphological features of buildups (Korolyuk 1960; shape of the laminae (plus microstructure in some
Krylov 1963) has won wide recognition. It is this system cases), and, for columnar forms, the presence or ab-
that meets the requirements of Proterozoic stromatolite sence of branching, its pattern, the general shape of the
biostratigraphy. Its basic categories are the form columns, and the type of their margins (plus micro-
( = form-species) and the group (=form-genus). In structure and/or lamina shape in some cases). The
addition, several suprageneric categories are in use and method of serial sectioning and graphic reconstruction
certain workers have distinguished subspecies or varie- proposed by Krylov (1963) provided the common basis
ties. for revealing these characteristics of columnar build-
 Proterozoic Stromatolite Taxonomy and Biostratigraphy 297

ups (see Walter 1976). In an attempt to express the dif- The discrimination and identification of form-spe-
ferences between taxa quantitatively, a morphometric cies (forms) in modern stromatolite classification is
approach to stromatolite systematics was proposed based either on: (1) a complex of three characteristics,
(Hofmann 1977, 1994) but has not gained wide recogni- minor variations in stromatolite lamination, morpho-
tion. logical details of the buildups, and the pattern of their
Different combinations of the above characteristics microstructure (texture), or (2) a combination of any
have been employed to specify all the valid stromatolite two of these characteristics, or (3) any one of them. In
genera known in the current literature and to verify the practice, the leading or single criterion for species
diagnoses and contents of some popular groups previ- identification is usually the microstructure, i.e. the dis-
ously established by using alternative criteria (e.g. Gym- tinctive features of the shape, construction, size, and
nosolen, Boxonia). The employment of the above charac- mutual arrangement of micritic or microsparitic com-
teristics provides a higher level of taxonomic resolution ponents of stromatolite laminae (Komar 1966). Accord-
for columnar stromatolites compared to stratiform and ingly, many species originally defined by morphologi-
nodular ones. At the same time, overestimation of the cal features alone have been revised according to the
diagnostic significance of minor morphological varia- details of their microstructure (e.g. Conophyton ressoti
tions of buildups of any kind has resulted in discrimina- Menchikoff, Boxonia gracilis Korolyuk, Linella ukka
tion of form-genera that are in fact environmentally de- Krylov).
pendent "ephemeral" modifications which should be re- The microstructural concept for the definition of
garded as junior synonyms of pre-existing taxa. Such formal stromatolite species initiated by Maslov (1960)
practice has locally been commonplace (e.g. Liang et al. and Semikhatov (1962) gained wide acceptance. Objec-
1984; Xing et al. 1985; Liang 1992) but is not widespread tions raised against this concept have been based on the
elsewhere because most specialists have reached a gen- obvious fact that the microstructure of any fossil stro-
eral understanding of the extent of the morphological matolite has been subjected to diagenetic alteration. To
attributes of buildups that are required for the identifi- date, however, the impact of these processes on stro-
cation of a valid stromatolite genus. matolite microstructure has been considered explicitly
It should be emphasized that any form-genus in the in only a few publications (Krylov 1963; Walter et al.
current stromatolite classification is defined by a com- 1988;. Fairchild et al. 1990 ). Nevertheless, stromatolite
bination of diagnostic features. Any single feature, no microstructures are usually considered to reflect as-
matter how spectacular and pronounced it may be, is pects of the stromatolite-forming microbial communi-
not sufficient for identification of a genus, although at- ties (Semikhatov et al. 1979; Walter 1983; Awramik 1991,
tempts to use incomplete sets of features for identifica- 1992; Walter et al. 1992, and references therein) and/or
tions of columnar stromatolites are known in the litera- secular change in global environments (Gretzinger
ture. Related misidentification of taxa at one time pro- 1990, 1994; Sochava and Podkovyrov 1992; Semikhatov
voked unsuccessful criticism of Proterozoic stromato- and Raaben 1994, 1996; Knoll and Semikhatov 1998).
lite stratigraphy (see Semikhatov 1978, 1991; Bertrand- Unfortunately, there is only one specific example of cor-
Sarfati and Walter 1981; Raaben 1981, for details). relation between microbiota composition, microstruc-
A particular problem in the designation of stromat- ture and stromatolite morphology: three vertically in-
olite form-genera is related to the current conception of tergrading varieties of Early Proterozoic silicified stro-
an abiogenic precipitated origin of some Precambrian matolites differing in both microstructure and gross
stromatolites, and especially of microstromatites morphology that yield three statistically different sets
(Asperiform or small digitate stromatolites, etc.; Grot- of microfossils (Awramik and Semikhatov 1979). It is
zinger and Read 1983; Hofmann and Jackson 1987; interesting to note that associated silicified microphy-
Gretzinger 1990, 1994; Lowie 1994; Gretzinger and tolites (Osagia spp.) contain a particular set of micro-
Rothmans 1996; Kah and Knoll1996; Knoll and Semik- fossils other than any reported in stromatolites (Ser-
hatov 1998, and references therein). However, micro- geev et al. 1998).
stromatites are stromatolites in a field sense, are shown In practice, discrimination between the overwhelm-
to be distinctive members of many regional stromato- ing majority (up to So - 85%) of Proterozoic stromato-
lite assemblages (Semikhatov 1978; Raaben 1980, 1991; lite form-species is based, entirely or predominantly,
Makarikhin and Kononova 1983; Grey 1984, 1994a,b; Li- on microstructure and these are precisely the species
ang et al. 1984), and demonstrate a particular time which are most useful in stratigraphy. Species defined
range which is supposed to reflect directional secular by minor morphological features are usually, but not
change in global environments. In addition, some of necessarily (e.g. Linella ukka Krylov and L. simica Kry-
the small digitate forms appear to be biophoric as well lov), confined to metamorphosed Proterozoic succes-
as biogenic (Hofmann and Jackson 1987; Raaben 1998). sions (e.g. Krylov and Perttunen 1978; Makarikhin and
Therefore, it would be of no purpose to exclude them Kononova 1983). However, a portion of microstructur-
from the traditional stromatolite system. ally defined species is based on poorly preserved or re-
298 M.A. Semikhatov, M.E. Raaben

crystallized material (e.g. Bertrand-Sarfati and Awra- Development of an integrated stromatolite classifi-
mik 1992). The occurrence of undiscerned synonyms cation attracted the attention of many researchers.
among them is very likely. Maslov (1960 ), returning to a restricted notion of ge-
Variations in the evaluation of the relative impor- neric-level stromatolite taxa, suggested uniting certain
tance of characteristics employed in the definition of morphological groups into morphological types within
stromatolite taxa of any rank are another source of syno- his non-traditional classification. Korolyuk (1960) was
nyms. Incomplete and polyglot descriptions of taxa also the first to formally distinguish, within the framework
contribute to this problem. However, our experience in of the traditional system, three types of stromatolites -
the study of Proterozoic stromatolites indicates that no columnar, stratiform and nodular. This subdivision
more than a fifth of the species and genera of stromato- gained wide acceptance and later was supplemented by
lites described within the framework of the current clas- columnar-stratiform and columnar-nodular types
sification should be considered as synonyms. (Krylov 1963, 1975) and microstromatites (Raaben
Stromatolite taxonomy is complicated by lack of cor- 1980 ). More detailed schemes for Proterozoic stromato-
relation between morphological and microstructural lite classification (Table 1) that initially concentrated on
characteristics - identical microstructures are known columnar forms were subsequently expanded to in-
in buildups of different morphology, and morphologi- clude all morphologies (Konyushkov 1978; Liang et a!.
cally similar structures assigned to a particular genus 1984, 1985; Raaben 1986; Raaben and Sinha 1989; Liang
can yield various microstructures that offer a means of 1992). The classifications by Liang et a!. and Raaben
species definition. Such relationships have been cited and Sinha are similar as far as family-level is con-
as a reason for considering morphological and micro- cerned, but differ in the interpretation of the diagnostic
structural characters as non-hierarchical but parallel features and content of higher taxa. But the fact that in-
(Hofmann 1977). In practice, however, microstructural formal categories equivalent to formal suprageneric
criteria dominate the definition of species, comprise a taxa are often explored in stromatolites research, and
part of some generic diagnoses based primarily on furthermore that they show distinctive secular trends
morphological features, but are not considered in re- in diversity (Semikhatov and Raaben 1996), suggests
gard to suprageneric taxa. that suprageneric classification has a good chance of
being appreciated in the near future. At present, the
2.3 grouping of stromatolites into a hierarchy of types,
Suprageneric Categories groups (form-genera), and forms (form-species) is
generally accepted.
The increasing number of described stromatolites of
lower taxonomic rank demanded the recognition of su- 2.4
prageneric categories to embrace large groupings Alternative Classifications
united by the most common features of morphology.
The largest groupings (e.g. stratiform, columnar, co- Two draft schemes for alternative classifications of Pre-
lumnar branching) have been in use for more than cambrian stromatolites were published in the last de-
three decades and represent integral components of cades by Vlasov (1977) and Komar (1989, and references
modern stromatolite systematics. Hierarchic ranking therein). Vlasov strove to present a natural system and
of all groupings suggests that they can be regarded as as a starting point established the Family Tyssageata-
suprageneric taxonomic categories of four ranks. Au- cea, containing subordinate genera and species. How-
thors of recent classifications of stromatolites have ar- ever, all ofVlasov's taxa were defined on the basis of the
bitrarily denominated them families, orders, classes, very characteristics used for the definition in the taxa
and subtypes (conferring the type rank to Stromatoli- of the traditional system and were incorporated in it
thi Pia; Konyushkov 1978; Raaben 1986; Raaben and (Raaben and Komar 1982). Komar (1989) proposed that
Sinha 1989), or families, subtypes, types, and super- stromatolite classification should be based consistently
types with no references to the formal status of the on microstructure, and in particular on the morphol-
main entity (Liang et a!. 1984, 1985; Xing et a!. 1985; Li- ogy and spatial arrangement of its major elements. He
ang 1992). Although the application of biological no- suggested the recognition of four hierarchic formal
menclature to the suprageneric classification of stro- taxa based on these criteria: supertype, type, genus and
matolites was specifically advocated by Konyushkov species. Komar described two supertypes, based on
(1978), such practice has met objections related to the major features of the above elements, and five types de-
origin of the buildups. Consequently, it was suggested fined by the element's shape, distribution and mode of
that informal categories- the morphotypes- should be arrangement. He also suggested that the two lowest-
used (Semikhatov and Raaben 1994, 1996) which corre- rank taxa should be based on the configuration, spac-
spond to the most distinctive entities among the two ing and size of the elements. Unfortunately, details of
superior formal taxa of Raaben and Sinha (Fig. 1). the classification have not been elaborated and the rele-
 Proterozoic Stromatolite Taxonomy and Biostratigraphy 299

13~\ 14
1~0.
7~ 9
~
~
~~
- ~~.
~

Fig. 1. Sketches of some representative Precambrian stromatolites morphotypes.1-5 Stratiform stromatolites: 1,2 Stratifera Koroliuk, 3 Tys-
sagaetes Vlasov, 4 Irregularia Koroliuk, 5 Malginella Komar and Semikhatov; 6-10 Nodular stromatolites: 6-8 Domical: 6 Paniscollenia Ko-
roliuk, 7 Colleniella Koroliuk, 8 Plicatina Raaben, 9,10 "Cabbage-like": 9 Cryptozoon (sensu Pia), 10 Cryptophyton Raaben and Komar; 11-13
Columnar non-branching stromatolites: 11 Colonnella Komar, 12 Conusel/a Golovanov, 13 Conophyton Maslov; 14 Yakutophyton Schapova-
lova; 15 Songjiella Cao eta!.; 16-19 Microstromatites: (scale bar 1 em) 16 Asperia Semikhatov, 17 Lochmecolumella Liang and Zhang, 18 Mini-
columella Raaben, 19 Vetella Krylov; 20-30 Columnar branching stromatolites: 20 Gymnosolen Steinmann, 21 Boxonia Koroliuk, 22-24 Tun-
gussida: 22 Tungussia Semikhatov, 23 Anabaria Komar, 24 Baicalia Krylov, 25 Kussiella Krylov, 26-30 Intricatida: 26 Kussoidella Semikha-
tov, 27 Sundia Butin, 28 Pilbaria Walter, 29 Confunda Semikhatov, 30 Kanpuria Raaben

vant publications should be regarded as a "declaration mulae derived from English descriptive morphological
of intent", rather than the presentation of a substantial terms. The "taxa" of this well-known scheme corre-
classification. spond to the largest suprageneric entities of the tradi-
Among a number of earlier alternative stromatolite tional classification.
classifications (see Maslov 1960; Hofmann 1969, for re- None of the alternative classifications of Precam-
view and bibliography) only the following two are of brian stromatolites has been particularly successful.
broad interest. For the moment, in its present state, the traditional sys-
Maslov (1960) disapproved of the use of binomial tem, exploring both morphological and microstruc-
nomenclature for stromatolites and proposed a new tural features, provides a means for communication
polynomial scheme for their classification and naming among workers.
to be used in stratigraphic and environmental analysis.
Each important morphological and microstructural 3
feature was given a Latin name, and the title of a taxon Stratigraphic Significance
was formed by listing several of these names. This led
to unwieldy terminology that was not a substitute for Attempts to use Precambrian stromatolites as a biostrati-
the description of a taxon. graphic tool initially proved very promising, and they are
An environmentally oriented scheme (Logan et al. still a significant means of correlation for Proterozoic, es-
1964) was developed on the basis of modern buildups, pecially Meso- and Neoproterozoic, successions. How-
but occasionally was applied to Precambrian forms as ever, the available data demonstrate that some important
well. Logan et al. emphasized variations and intergra- limitations need to be taken into account.
dations in stromatolite morphology and proposed a The first empirical data underlying the current in-
system of symbolic Latin letter designations and for- ference about the stratigraphic importance of Protero-
Table 1. Successive stages in the development of the traditional stromatolite classification system ""00
Pia 1923 Genus Archeozoon Gymnosolen Weedia Collenia Cryptozoon
Maslov 1960 Morphotype Conophyton Collenia ;!::
Supergr. C. columnaris C. undosa ?>
Koroluk Type Subtype Columnar Stratiform Nodular s"'
1960
~
Krylov 1963 Type Columnar Strati co- Stratiform (Columnar)
"'
~
lumn. ;!::
Subtype Branching Nodular ~
;;d
Raaben Type Columnar Stratiform Nodular
1964
~
Non-branching Actively Passively branching g
1969 branch.
Categorgy Conophytonida Gymnosole- Kussiellida
nida
Tungussida
Komar 1966 Type Columnar Non-columnar
Subtype Non-branching Branching Stratiform Nodular
Actively Passively branch.
branching
Krylov 1975 Type Columnar Stratico- Stratiform Column Nodular
lumn. nodul.
(unnamed) Branching
(I) (2) (3)
Koniushkov Subtype Columnar (Colonnelithi) Stratiform (Stratiferithi) Nodular
1978 Class Non-branching Branching (Gymnosolenitha) Strati co- Stratiferitha
( Colonnelitha) lumn.
Order Colonnelida Gymnosolenida Kussi- Omach- Stratiferida
ellida tenida
Family Alveolidae Gymnosolenidae Kussiel- Omach- Irregularidae
Colonnelidae Tungussida lidae tenidae Stratiferidae
Conophytonidae
Raaben et Subtype Columnar (Columnithi) Non-columnar (Compactith i) Microstroma-
Sinha 1989 Class Non-branching Branching (Ramificantha) Tabulitha (Stratiform) Pienostroma (Nodular) tithi
Order Con ophytonida Intricatida Acrescentida Kussiellida Anaglip- Decumbenta Cupolida Cryptiida Minicolumel-
(Irregularly (Actively (Passively branching) honida (Domelike) !ida
branching) branching)
Family Euconophy- Prokussielli- Gymnosole- Kussiellidae Stratiferidae Malginelli- Tinnidae Cryptophy- Minicolumel-
tonidae dae nidae Omachtenidae Tyssagaet- dae Confluentidae tonidae lidae
Ephyaltidae Discorsiidae Tungussidae idae Bulboidea Pseudogym-
Kanpiridae Alterniidae nosolenidae
lllietidae
 Proterozoic Stromatolite Taxonomy and Biostratigraphy 301

zoic stromatolites were obtained in the 1960s in the platforms, together with the environmental specializa-
course of the study of the Riphean and Vendian build- tion of some distinctive morphotypes (e.g. Serebrya-
ups in northern Eurasia (see Raaben 1969; Semikhatov kov 1975; Grey and Thorne 1985; Grotzinger 1990; Zhu
1976, for review and bibliography). During these years and Chen 1992), predetermines the discrete distribu-
an advanced version of stromatolite classification was tion of these biolites and of their assemblages in the
presented (see above), and reliable data on the time- sedimentary successions. In this respect, the Uralian
and-space distribution of the Late Proterozoic taxa stratotype of the Riphean is no exception, since stro-
were obtained for the first time. It was revealed that the matolites are present here only in the middle and/or
Riphean and Vendian sections from several widely sep- upper parts of the unconformity-bounded type units of
arated regions of Siberia and the Urals yield directional the Lower, Middle, and Upper Riphean erathems (Kry-
successions of taxa of columnar stromatolites, and that lov 1975; Raaben and Komar 1982). Their lower bound-
these successions, in the most complete sections, fall aries are traditionally located at unconformities at the
into four, time-specific, taxonomically distinct com- bases of siliciclastic stromatolite-free formations that
plexes laterally linked by a number of common genera constitute the lower parts of these type units. Attempts
and some common species. In addition, K-Ar glauco- to substitute Siberian sections, containing more pro-
nite ages of the comparable complexes in different re- lific stromatolites, for the Uralian ones in stromatolite
gions were shown to be similar (Keller et al. 1960; Se- definition of the boundaries (e.g. Semikhatov 1974)
mikhatov 1962; Krylov 1963; Raaben 1964; Komar 1966). cannot be regarded as a correct approach for two rea-
Before long, these results were supported by data from sons. Firstly, the Siberian sections also show discrete
other continents (e.g. Bertrand-Sarfati 1972; Walter time distribution of stromatolite assemblages. Sec-
1972, 1976; Preiss 1972, 1973, 1977; Bertrand-Sarfati and ondly, such substitution, quite trivial in the Phanero-
Walter 1981, and references therein), and by the record zoic, is fraught with serious misleading consequences if
of non-columnar morphotypes (e.g. Komar 1966). applied to the Proterozoic. This is due to the limited
The accumulated data suggested that stromatolites possibility for precise long-distance correlation in this
are an effective tool for both general four-fold subdivi- part of the geological record and to strong lateral varia-
sion and interregional correlation of the Late Protero- tions in the taxonomic composition of stromatolite as-
zoic (e.g. Keller et al. 1960; Semikhatov 1962, 1974; Kry- semblages.
lov 1963, 1975, 1985; Cloud and Semikhatov 1969; Raa- In other words, researchers in the 1960s -1980s over-
ben 1969, 1975; Bertrand-Sarfati 1972; Walter 1972, 1976; looked the obvious fact that Riphean (as well as any
Preiss 1977). Occurrence of specific stromatolite com- other) stromatolites are unsuitable for defining chrono-
plexes within the type section of each previously recog- stratigraphic boundaries, and, hence they cannot serve
nized Riphean erathem (the Lower, Middle and Upper as a tool for subdivision of the Proterozoic. This be-
Riphean) provided, for the first time, paleontological came clear later, when attention was focused on the ap-
characteristics of these widely used units and created plication of chronostratigraphic requirements to Pro-
the impression that they were paleontologically defined terozoic stratigraphy (Semikhatov 1991, 1993, 1995).
subdivisions with distinct boundaries. This view was However, stromatolites provide unique, essentially
facilitated by the fact that stromatolite workers in the paleontological, characterization of units which have
1960s - 1970s failed to discriminate between two chro- been defined by other methods. They contribute to
nostratigraphic procedures - subdivision and correla- both the establishment of the paleontological charac-
tion. terization of the units, and their correlation in sepa-
Although initial results looked promising, subse- rated sections. The time-and-space distribution of such
quent studies revealed a number of problems in Prote- units is defined by the time range and lateral persis-
rozoic stromatolite biostratigraphy related to taxon- tence of successive stromatolite complexes.
omy, discrete distribution of assemblages in the succes- The available data demonstrate that the most dis-
sions, provincialism of time-dependent taxonomically tinct change in the Proterozoic stromatolite succession
distinct complexes, and variability in the time ranges of occurred across the Paleo-Mesoproterozoic transition.
taxa at the interregional scale. An appreciation of these Although at one time b elieved to be similar to the Late
problems and of the requirements of chronostratigra- Riphean complex (Hofmann 1977, and references
phy are the basis for the current evaluation of the strati- therein), the Paleoproterozoic stromatolite complex
graphic potential of Precambrian stromatolites (Se- was shown to be dominated by genera and species un-
mikhatov 1991, 1995; Raaben 1997). known in the Riphean and Vendian. Most Paleoprote-
In evaluating the applicability of stromatolites for rozoic forms "of Upper Riphean aspect", which sug-
general subdivision of the Proterozoic, i.e., for defining gested the taxonomic similarity of Paleo- and Neopro-
the boundaries of chronostratigraphic units, several terozoic stromatolites, were assigned to peculiar taxa at
factors have to be taken into account. The confinement both specific and generic levels (Semikhatov 1978; Raa-
of the bulk of Proterozoic stromatolites to carbonate ben 1980, 1991, 1997; Makarikhin and Kononova 1983;
302 M.A. Semikhatov, M.E. Raaben

Grey 1984, 1994a,b; Liang eta!. 1984; Xing et a!. 198s; Se- Variability of the time range of taxa at interregional
mikhatov and Raaben 1994, 1996). But despite its dis- and, especially, interprovincial scale presents addi-
tinctive taxonomic composition and very large time tional problems in the stratigraphic employment of
range, the Paleoproterozoic stromatolite complex can Proterozoic stromatolites. For instance, Conophyton cy-
only be subdivided into subordinate, stratigraphically lindricus Maslov is confined only to the lower Upper Ri-
useful taxonomic entities on a regional scale (e.g. Grey phean in Eastern Siberia, but in the integrated north
1984; Liang et al. 1984). As for the Meso- and Neoprote- Eurasian scale it is distributed from Lower Riphean to
rozoic (Riphean and Vendian) stromatolite succession, lower Upper Riphean. Similarly, Kussiella kussiensis
it falls into four intercontinentally recognizable assem- Krylov is confined to the lower Lower Riphean in the
blages of taxa which provide the paleontological charac- Urals, northern Siberia, northern Australia, and India
teristics for the Lower, Middle, and Upper Riphean and but is reported from the Lower and lower Middle Ri-
Vendian (e.g. Krylov 197s; Semikhatov 1976, 1991; Ber- phean in SE Siberia (Uchur-Maya Region), and Linella
trand-Sarfati and Walter 1981, and references therein). simica Krylov occurs in the lower upper Riphean in
The datings of the lower boundaries of these units are eastern China, in the uppermost part of the Upper Ri-
considered to be 16so ± so, 13SO ± 20, 1000 ± so 1, and phean in the Urals, and in the Vendian-Cambrian
6so ± 20 Ma (Semikhatov 1993 and references therein). boundary beds in the Uchur-Maya Region.
A survey of available data on the distribution of strom at- Judging by independent correlation criteria, not
olite genera and species demonstrates that in certain only particular taxa but also some associations of taxa
vast regions (e.g. northern Eurasia, China) some infor- can change their time range when passing from one
mal subdivisions that are subordinate to the three Ri- province to another.
phean erathems also have specific stromatolite charac- This is not to say that all stromatolite taxa are wide-
teristics (Krylov 1975, 198s; Raaben 1969, 197s; Liang et ranging and time-transgressive at the interprovincial
al. 1984, 198s; Semikhatov and Raaben 1994, 1996). scale. Some characteristic members of regional assem-
It is significant that time-dependent successions of blages possess relatively short (by Proterozoic stan-
Proterozoic stromatolites are governed by the secular dards) time ranges, both at regional and interregional
change of time-restricted taxa defined by different scales, and can be used for long-distance correlation.
combinations of selected attributes. The notion that at To take the most convincing examples, Baicalia lacera
one time was popular, of directional development Semikhatov, which stands out because of its very dis-
("evolution") of their real attributes (e.g. Krylov 1963, tinctive microstructure (Knoll and Semikhatov 1998),
197s), lacks support from recent data. Consequently, is restricted to lower Upper Riphean sections in Siberia,
Proterozoic stromatolite stratigraphy remains strictly northern Africa, north-western Canada, and south-
empirical. western Alaska, while ]urusania nizvensis Raaben was
Important constraints on the stratigraphic value of reported from the lower or middle part of the Upper Ri-
Proterozoic stromatolites in long-distance correlation phean in the Urals, eastern China and South Australia
are imposed by distinct provincialism in time-depen- (see Semikhatov and Raaben 1994, 1996 for bibliogra-
dent complexes (Semikhatov 1980, 1985, 1991; Golove- phy). Unfortunately, the number of such relatively
nok 1984). The degree of provincialism is evident from short-ranging but laterally widespread (interprovin-
the fact that among all the taxa present in five, Middle- cial) taxa is very limited.
Late Riphean, northern Eurasian and African stromat- Taken together, these considerations indicate that
olite provinces, only 12% of species and 52% of genera stromatolite-based interprovincial correlations should
are known to extend beyond the boundaries of at least be based on similarities in the secular successions of
one province. A survey of the global data shows that a particular stromatolite assemblages, rather than on oc-
high degree of endemism of Proterozoic (and especially currence of individual common taxa, and wherever
pre-Riphean) stromatolites is the rule rather than the possible must be supported by radiometric and/or che-
exception - only about 8.s% of species and 31% of gen- mostratigraphic data. Such correlations allow the trac-
era occur in common in two or more super-regions ing, at the interprovincial scale, of Paleoproterozoic,
such as northern Eurasia, China, India, Africa, North Lower, Middle, and Upper Riphean and Vendian de-
America, and Australia (Semikhatov and Raaben 1996). posits, and, between some provinces, of particular
Consideration of the so-far-unrecognized synonyms parts of Riphean erathems.
will somewhat change the above figures, but is unlikely The effectiveness of stromatolites in the intrabasinal
to affect the general situation with regards to the high correlation of Proterozoic successions is much higher.
endemism of Proterozoic stromatolites. It is convincingly demonstrated by many Late Protero-
zoic (e.g. Semikhatov 1962; Raaben 1964, 1975; Komar
1966; Bertrand-Sarfati 1972; Semikhatov and Serebrya-
1 New isotope-geochronological data obtained in Siberia demon-
strate that the Middle-Upper Riphean boundery should be dated at kov 1983; Liang et al. 1984; Raaben eta!. 1995) and Early
1050 ± 20 Ma Proterozoic examples (e.g. Makarikhin and Kononova
 Proterozoic Stromatolite Taxonomy and Biostratigraphy 303

Fig. 2. Vertical distribution of stromatolite taxa in the Riphean and Vendian of the Uchur-Maya Region and its geological setting (inset
map; modified from Semikhatov and Serebryakov 1983). 1,2 Dominant rock composition: 1 carbonate and siliciclastic-carbonate, 2 silicic-
lastic, 3 principal unconformities, 4 vertical distribution of dominant (a) and subordinate (b) taxa in assemblages; 5-9 on the inset map:
5 pre-Riphean rocks, 6 Riphean and Vendian: 7 Phanerozoic, 8 principal thrusts and upthrusts, 9 other faults. Formations: Gn Gonam; Om
Omakhta; En Ennin; Tr, Trekhgorka; Dm Dim; Tl Talyn; Sv Svetly; Tt Totta; Mig Malgina; Tsp Tsipanda, Nr Neryuen; Ign Ignikan; Kn Kan-
dyk; Uk Ust'kirba; Am Aim; Ujd Ust'-Yudoma

1983; Liang et al. 1984 1985; Grey 1984, 1985), amongst and most genera) have definite, though in part very
which the Riphean and Vendian stratigraphically com- large, time ranges at the interregional scale.
plete and stromatolite-rich succession of the Uchur- Secular change in taxa defined the dynamics of the
Maya Region can be considered a model (Fig. 2). It global taxonomic diversity of Proterozoic stromato-
demonstrates the occurrence of a unique vertical lites. Analysis of theses dynamics at specific and ge-
change in stromatolite taxa and their grouping into def- neric levels in an environmental context, and with a rel-
inite assemblages of various diversity which differ from atively high time-resolution (Semikhatov and Raaben
one another at both the generic and, especially, at the 1994, 1996), clarified the previously established pat-
specific levels. Each assemblage characterizes a certain terns (Walter and Heys 1985; Awramik 1992) and re-
interval of the section equal to, or smaller than, the re- vealed that the main variations in stromatolite diversity
gional lithostratigraphic units. Assemblages are usu- have been linked by direct relationships and feed-back
ally delimited by siliciclastic horizons and/or uncon- mechanisms to abiotic, biologically induced, and biotic
formities; only two assemblages intergrade in the sec- events. The most important among these were global
tion. As to lateral distribution, some assemblages tectonics (partly reflected in the variations in mantle
maintain their taxonomic composition throughout the fluxes and continental run-off into the World ocean, as
whole region while others show rapid changes over recorded in the Sr-isotopic composition of sea water),
short distances that are not necessarily in concert with significant climatic variations and related glacioeusta-
lithological variations in the enclosing rocks (see Se- tic sea-level fluctuations, directional changes in pC0 2,
mikhatov and Serebryakov 1983). the degree of carbonate oversaturation in surface sea
The Uchur-Maya section, together with other exam- water and related evolution of carbonate sedimenta-
ples, shows that although the distribution of a regional tion, secular variations in the production and burial of
stromatolite assemblage in time and space was clearly organic and carbonate carbon, and, last but not least,
affected by environment, environmental pressure is not evolutionary changes in the Riphean and Vendian bi-
the main, and even less the only, factor governing the ota. The last link is evident from the chronological co-
taxonomic composition and time range of a particular incidence between important variations in Neoprote-
assemblage, since an overwhelming majority of Prote- rozoic stromatolite diversity and principal biological
rozoic stromatolite lower-rank taxa (almost all species events (see Knoll and Sergeev 1995; Fedonkin 1996, and
304 M.A. Semikhatov, M.E. Raaben

references therein). It may, therefore, be deduced that of any assemblage was determined by its time-and-
the very existence of time-restricted stromatolite taxa space position with regard to the time range of rele-
and their assemblages was defined by the evolution of vant stromatolite taxa and their preferred environ-
global environments and, partly, of microbial commu- mental settings.
nities, on the early Earth. The documented relation- 5. Interprovincial correlations by means of stromato-
ships explain an obvious contradiction between evolu- lites are oflower reliability and time-resolution than
tionary conservatism of stromatolite-forming micro- intraprovincial ones and should be constrained by
organisms and the empirically established secular independent methods of long-distance correlation.
change of Proterozoic stromatolite taxa. In the present 6. Precambrian stromatolites are unique objects in
state of the art, these relationships confirm Maslov's Earth history, since the predominance of microbes
(1959) surmise that the stratigraphic potential of Pre- in ecosystems, as well as the global environments
cambrian stromatolites was defined by the evolution of themselves and the scale of their evolution recorded
both environmental and biological factors relevant to in the Archean and Proterozoic, had no counterparts
the construction and facies restrictions of these bia- in the Phanerozoic.
lites.
Acknowledgements. We sincerely thank S.M. Awramik, K. Grey,
4 and the late V. Komar for fruitful discussions of the problems con-
sidered in this paper, and gratefully acknowledge the constructive
Conclusions comments of the reviewers of the manuscript. This work was sup-
ported by the Russian Foundation for Basic Research, Project
1. The traditional stromatolite system has proved to be 96-05-64329.
the most useful in the study of Proterozoic buildups.
A number of stromatolite classifications published
during the past decades represent successive stages
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