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Evolution of trilobite biofacies in Cambrian basins of the Siberian platform

Evolution of trilobite biofacies in Cambrian basins of the Siberian platform

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Journal of Paleontology: Evolution of trilobite biofacies in Cambri...http://findarticles.com/p/articles/mi_qa3790/is_200011/ai_n892...1 of 68/12/2008 12:10 AM
FindArticles>Journal of Paleontology >Nov 2000> Article> Print friendly 
Evolution of trilobite biofacies in Cambrian basins of the Siberian platform
Pegel, Tatyana V  ABSTRACT-Cambrian biotic zonation on the Siberian Platform reflects differentiation of the depositional environments (inner shelf, outer sheland open basin). The combination of the chart of trilobite biofacies replacement and the curve of sea-level fluctuations shows that trilobite biofacies replacement occurs as a rule at times of sign reversal and distinct change in the rates of sea-level rise or fall. The boundaries of majorSiberian platform Cambrian chronostratigraphic units, such a stages and series, frequently coincide with the boundaries of biofacies instratigraphic succession related to sea-level fluctuations. If these fluctuations are gradual and restricted, then the boundaries of the Cambrianstages and series cannot be isochronous levels at a global scale. The known levels for intercontinental correlation on the Siberian Platform include boundaries of the adjacent Triplagnostus gibbus and Tomagnostus fissus Zones from the uppermost Amganian Stage (Middle Cambrian) and theGlyptagnostus stolidotus and Glyptagnostus reticulatus Zones of the lower Upper Cambrian. Both levels correspond to boundaries betweenhighstands and lowstands on the Siberian Platform and appear to serve as boundaries of high rank. Evolution of the trilobite biofacies zonation isillustrated by genera typical for each of the various Cambrian paleogeographic environments on the Siberian Platform.INTRODUCTIONTHE BIOLOGICAL history of the Cambrian basins on the Siberian Platform has been traditionally studied in parallel with the lithostratigraphy, but to a great extent separately, in the context of biostratigraphy (i.e., in time sequence) and biofacies or paleoecology (i.e., in areal extent). As aresult of reconstruction of environmental and depositional conditions for the constantly changing paleobasins, the concept of sequencestratigraphy makes it possible to consider organism assemblages in three-dimensional space for various time intervals. It allows one to reveal andtrace cause-and-effect relations of biotic and abiotic components of the evolving paleobasins.The Cambrian biotic zonation on the Siberian Platform (Fig. 1) reflects differentiation of depositional environments initiated in Vendian time andpreserved during the entire Cambrian (Pisarchik et al., 1975; Melnikov et al., 1989; Sukhov, 1997). On the trilobite biofacies maps (Figs. 2-4)the pattern of depositional environments is that proposed by Sukhov (1997). Correlation between the spatial distribution of trilobite associationsand depositional environments shows that environmental units of high rank (inner and outer shelves, basin slope) are characterized by specifictrilobite assemblages (biofacies) during time intervals equivalent to ages, epochs or even longer. These assemblages are characterized by one or afew taxa generally of high rank (family, superfamily, order) depending on the quality, diversity, frequency and distribution of the named taxa.The leading role of lower-rank taxa (genera and species) is confined, as a rule, to correlation within contemporaneous environments exhibitingfacies similarity. In some facies regions, side-by-side with a number of relatively short-lived biofacies, there may be a long-lived biofacies (heremacrobiofacies) usually characterized by a high-rank taxon which was also present in the short-lived biofacies.Biotic development is a reflection of the history of geographic, hydrodynamic and tectonic changes to the paleobasins. Among them, eustaticsea-level fluctuations are of primary importance. The combination of the Cambrian chart of trilobite biofacies zonation on the Siberian Platformand the curve of sea-level fluctuations constructed by Sukhov (Pegel and Sukhov, 1996) shows that trilobite biofacies replacement occurs as arule at times of sign reversal and distinct change in the rates of sea-level rise and fall (Fig. 1). Shelf and particularly inner-shelf assemblages, where biofacies replacement was sometimes more frequent than in other paleogeographic zones, are most sensitive to associated variations inphysical and chemical parameters of the environment (temperature, oxygen and hydrogen sulfide saturation, illumination, flow rate, etc.).The Cambrian trilobite fauna on the Siberian Platform during its long evolution retained specific features formed in certain environments.Trilobite associations on the slopes of deep-water open basins, occupying the eastern and occasionally northwestern parts of the Siberian Platform,are characterized by high diversity in sizes of individuals and in taxonomic composition. Genera and species having global distribution weresignificant components, mainly in assemblages from the lower parts of the slopes. These were primarily miomeroids and to a lesser extentpolymeroids. The inner shelf, covering the central and southwestern Siberian Platform, represented a semi-closed relatively shallow water basinoccasionally with high salinity. In trilobite assemblages there, the index taxa are frequently genera. The faunas are typically endemic,individuals are commonly small but abundant, and taxonomic diversity is low. Fossiliferous beds are rare and most commonly found in sectionsadjacent to the outer shelf. The inner shelf and deep-water open basins were separated by a rather narrow band of outer shelf shallow carbonateplatforms; reef formation on them was recurrently active. This zone was most favorable for trilobites. Fossil complexes found there are highly diverse in composition so that index taxa are difficult to define. These taxa are known primarily from trilobite associations of the reef slope only.The trilobite fauna of the outer shelf is characterized by individuals of various sizes and by significant endemism. The boundary areas of themajor depositional environments are characterized by trilobite associations combining elements of communities from neighboring facies zones.This is especially noticeable between the outer shelf and upper slope of the open basin. Thus, the trilobite faunas of these depositionalenvironments are frequently characterized by common index taxa.The oldest trilobites on the Siberian Platform are polymeroids of the Fallotaspid Biofacies (Profallotaspis, Repinaella) inhabiting the outer shelf reef and bank zone in the first half of the Atdabanian (Early Cambrian) (Figs. 2.2, 7). The first miomeroids of the Pagetiellus Biofacies, initiatingthe Pagetiid Macrobiofacies, appeared in the deep open shelf in later Atdabanian time. In the later half of the Atdabanian, representatives of theFallotaspidoidea (mainly Judomia) and Pagetiidae (principally Pagetiellus and to a lesser extent Hebediscus and Triangulaspis) occupy bothecological niches and constitute a single Fallotaspid-Pagetiid Biofacies (Figs. 7, 10).
Journal of Paleontology: Evolution of trilobite biofacies in Cambri...http://findarticles.com/p/articles/mi_qa3790/is_200011/ai_n892...2 of 68/12/2008 12:10 AM
 With rise in sea level in Botomian time, the Fallotaspid Biofacies of the outer shelf was replaced by numerous trilobite assemblages represented by a variety of taxa, where the index taxon is difficult to determine. The assemblages include Dorypygidae (Kootenia), Dinesidae (Erbiella, Erbiopsis,Rondocephalus, Proerbia), Dolichometopidae (Poliellina), Ptychopariidae (Binodaspis), Namanoiidae (Inouyina, Chondrinouyina),Emmrichellidae (Sanaschtykgolia), Aldonaiidae (Aldonaia), Granularaspidae (Granularaspis), Judomiidae (Judomiella), Pagetiidae (Pagetiellus,Hebediscus, Neopagetina, Neocobboldia) and many others. These assemblages are here provisionally named the Dorypygid etc. Biofacies. In thereef slope (the external part of the outer shelf), Protolenidae (diverse species of Bergeroniellus) were dominant (Figs. 2.3, 7, 10).In Botomian time, the slope/basin environments were most favorable for the Protolenidae (primarily Bergeroniellus) and Pagetiidae (Pagetiellusand Neopagetina). In Early Botomian time, the Calodiscus Biofacies was associated with black shales of the deep water zone (Figures 2.3, 10).On the Early Cambrian inner shelf, more frequent substitutions of biofacies occurred that had nothing in common with faunas of the outer shelf and open sea. At the end of Atdabanian time, the Neoredlichiid-Malykaniid Biofacies (Elganellus, Buliaspis, Malykania) was widely developed(Figs. 2.2, 5). This was replaced by the Jakutiid Macrobiofacies (Bathyuriscellus, Jakutus) in Botomian time. Simultaneously at the beginning of Botomian time Redlichiidae (Tungusella) became of considerable importance in the inner shelf faunas. During a strong rise in sea level inBotomian time a brief expansion of Protolenidae (Bergeroniaspis) from the open sea took place and they also became characteristic elements of trilobite assemblages in the inner shelf region (Figs. 2.3, 5). A significant drop in sea level at the beginning of Toyonian time resulted in a strong faunal change in the inner shelf region, reflected by replacement of the biofacies index taxa at both generic and family levels (Pseudoeteraspis and Parapoliella in Early Toyonian, and Namanoia inLate Toyonian time). However, this event caused no distinct changes in biofacies of the outer shelf and open sea regions, even at the family level(Figs. 3.1, 5, 7, 10).The Lower-Middle Cambrian boundary marks a major change of taxonomic composition of the trilobite faunas over the entire Siberian Platform.This event coincides in time with the commencement of active sea-level rise and renewed reef building. Distinct biofacies differentiation not only of the inner and outer shelf faunas but of the shelf/basin ones took place as well (Fig. 3.2). Nevertheless, the faunal relations across the entireSiberian Platform remained rather intimate: characteristic elements of shelf communities (Pseudanomocarina, Chondranomocare,Chondragraulos) are relatively common in basinal biofacies, and characteristic representatives of the basinal facies (agnostoids and eodiscids)occur in shelf biofacies.In Amganian time, the outer shelf is characterized by the Anomocarid-Dorypygid Biofacies where Anomocaridae are represented mainly by Chondranomocare; Pseudanomocarina; Kootenia and Olenoides were most abundant among the Dorypygidae; the Dinesidae were represented by Dinesus; and the Emnmrichellidae by Chondragraulos (Fig. 8).Throughout Amganian time, the open basin was characterized by the Paradoxidid Biofacies (Anabaraspis, Paradoxides). In Early Amganian timethere was a period when the paradoxidids were less abundant and less frequent in trilobite assemblages than Oryctocephalidae (Oryctocara,Oryctocephalops, Oryctocephalites, Cheiruroides) which thus characterized a short-lived Orycrtocephalid Biofacies. In Late Amganian time, thetrilobite assemblages of the open sea show predominance not only of Paradoxididae, but of agnostoids as well. The latter form the AgnostidMacrobiofacies, which retains its importance in the open sea environments, mainly on the lower slope, until nearly the end of the Cambrian (Fig.10).The inner shelf environments in Amganian time are characterized by the Asaphiscid Biofacies (Proasaphiscus, Deltocephalus, Itcheriella) (Fig.5).It should be stressed that in the unified Cambrian charts for the Siberian Platform (Krasnov et al., 1983; Astashkin et al., 1991) officially adoptedin Russia, the Lower-Middle Cambrian boundary in the Yudoma-Olenek region, which corresponds to the open basin, is placed not at theemergence of the first numerous paradoxidids, represented by Anabaraspis, as suggested by Savitsky and Shabanov (Savitsky et al., 1972;Egorova et al., 1976) but somewhat higher, at the base of the Oryctocara Zone. However, lithofacies analysis shows that it is the base of the Anabaraspis Zone rather than the base of the Oryctocara Zone that coincides with the level reflecting regional events, i.e., onset of a strongsea-level rise and renewed reef building. A Lower-Middle Cambrian boundary at the base of the Anabaraspis Zone of the Yudoma-Olenek and Anabar-Sinyaya facies regions perhaps corresponds to the top of the Namanoia Zone in the Turukhan-IrkutskOlekma facies region, which is theofficial top of the Russian Lower Cambrian; but it is also possible that some upper part of the Namanoia Zone corresponds to the Anabaraspis Zoneas reflected in official Russian charts (Krasnov et al., 1983). At the very end of Amganian time, a sharp drop of sea level resulted in a large part of the inner shelf, until Middle Mayan time, becoming a low supratidal plain (Fig. 4.1). Trilobites reappeared there only when the region was re-submerged. In Late Mayan time, two endemic genera,Markhaspis and Kuraspis became its inhabitants (Fig. 5). The trilobite associations found are extremely uniform in composition and endemic,sometimes represented by only one genus with strong intraspecific variability and numerous small individuals. The trilobite fauna of thetransitional zone between the inner and outer shelves, inhabited at that time by very rare representatives of Ritella and some other genera, hassimilar characteristics (Fig. 6). An Early Mayan lowstand and active sediment filling of the deep-water basin on the eastern Siberian Platform drastically reduced the diversity of the trilobite faunas of the outer shelf and promoted their displacement toward the open basin region. Thus, the Anomocaridae became a biofacies index taxon along with Dorypygidae, not only for the outer shelf but also for the upper slope of the basin, replacing there the Amganian
Journal of Paleontology: Evolution of trilobite biofacies in Cambri...http://findarticles.com/p/articles/mi_qa3790/is_200011/ai_n892...3 of 68/12/2008 12:10 AM
Paradoxidid Biofacies. The trilobite assemblages are characteristically represented by such genera as Irinia, Harataspis and Pseudanomocarina,and to a lesser extent by Olenoides and Basocephalus; Semicyclocephalus is also rather characteristic (Fig. 8).The Middle Mayan trilobite fauna of the outer shelf is still unknown. However, in the transitional zone from the external margin of the carbonateplatform to the upper slope, the trilobite associations are taxonomically diverse and the index taxa for the biofacies remain the Anomocaridae(Harataspis, Pseudanomocarina, and less commonly Anomocarioides and Anomocarina) and Dorypygidae (Dorypygoides, Rabutina). Anomocarellidae such as Bonneterrina, Sacha, Usoviana, Aldanaspis, and new genera of the Anomocarellidae are also characteristic, andagnostoid trilobites appear. At the end of Mayan time, the outer shelf Anomocarid-Dorypygid Biofacies was replaced by the Plethopeltid Biofacies. Distinctive elementsinclude Koldinia, Koldiniella, Maiaspis, Buttsia, Seletella, Onchonotellus, Bonneterrina, Sacha, and less commonly Schoriecare and rare Agnostus? aff. simplexifonis Rozova (Fig. 8).In Early Mayan time, the upper slope is the region of the Anomocarid-Agnostoid Biofacies where various agnostoids (Goniagnostus,Dolichagnostus, Triplagnostus, Megagnostus) prevailed. In Late Mayan time, anomocarids (Anomocarioides and Schoriecare) were most typical.Rather frequent additional elements include Anomocarellidae (Liopelta and other genera) in Early Mayan time and Plethopeltidae(Plethopeltoides, Koldinia, Koldiniella), Bonneterrina and Aldanaspis in Late Mayan time (Fig. 11). Throughout Mayan time diverse agnostoids were primary constituents of the lower slope trilobite assemblages.The transgression of the second half of Mayan time, and continuing into the beginning of the Late Cambrian, was gradual and accompanied by small fluctuations of sea level. Trilobite associations showed gradual evolutionary changes, but general continuity in composition from Middleinto early Late Cambrian time, as well as gradual faunal transitions between different environmental regions. Thus, members of thePlethopeltidae, the index biofacies taxon for the outer shelf, expanded their ranges to become characteristic taxa of biofacies in adjacent parts of the inner shelf and upper slope of the slope-basin region (Fig. 4.2).The Upper Cambrian faunas of the western part of the inner shelf known at present represent only the Ayusokkanian and most of the Sakianstages. Only representatives of Kuraspis, the index taxon of the Kuraspis Biofacies, inhabited this area of the paleobasin, except at the extremesouthern part of the Siberian Platform where it was accompanied by Verkholenella. The eastern part of the inner shelf, reflecting the proximity of organic buildups of the outer shelf, also has frequent representatives of the Plethopeltidae (Koldinia) and the less common genera Markhaspisand Letniites (Fig. 6). The trilobite associations in this part of the basin are more diverse taxonomically, though poor in individual quantity.Faciura, Pesaiella, Bolaspidina, Acidaspidina, Raashellina and Nordia may also be present. Generally, the trilobite fauna in the marginal part of the inner shelf consists of elements of the more diverse trilobite assemblages of the outer shelf, mainly its inner part. These two adjacentpaleogeographic regions constitute the area of the Plethopeltid Macrobiofacies during Ayusokkanian and most of Sakian time.In Ayusokkanian time, within the Plethopeltid Macrobiofacies, the Nganasanella Biofacies is prominent in the area of the outer shelf. In additionto Nganasanella, genera such as Koldinia, Koldiniella, Acrocephalella, Pauciella, Acidaspidella, Pedinocephalina, Pesaiella, Theodenisia, Seletella,Munija, Schoriecare, Catuniella, Buttsia, and Ritella are characteristic elements of the biofacies (Fig. 9). At the end of Ayusokkanian and the beginning of Sakian time (Fig. 1), sea level rise was replaced by gradual fall. It is most evidently, but notdrastically, reflected in the fauna of the outer shelf where replacement of less important taxa took place. At the beginning of Sakian timePterocephaliidae appear and become a characteristic element of the Ceratopygid Biofacies on the slope, and the Pterocephaliid-ParabolinidBiofacies replaces the outer shelf Nganasanella Biofacies of the Plethopeltid Macrobiofacies. Distinctive taxa of the Pterocephaliid-ParabolinidBiofacies include Maduiya, Idahoia, Pesaiella, Faciura, Nganasanella, Koldinia, Plethopeltoides and Cyclognathina (Fig. 9). At the end of Sakian time, as the sea level continued its gradual fall accompanied by small fluctuations, there was a pronounced depletion of thetrilobite fauna of the outer shelf. At this time, and into Aksayan time, the Pterocephaliid biofacies (Amorphella, Olentella, and Monosulcatinaaccompanied by Yurakia, Polyariella, Ketyna, Dolgeuloma and Kaninia) became established on the outer shelf and the adjacent part of the innershelf (Fig. 9).The fauna of the slope/basin, primarily the lower slope, was less affected by sea-level fluctuations. During most of Middle Cambrian and LateCambrian time the Agnostoid Macrobiofacies dominated this region. More frequent replacement of biofacies occurred on the upper slope, but eventhose changes were not as obviously connected to sea level fluctuations as the changes in the outer shelf region.During Ayusokkanian time and most of Sakian time, the transitional area between the outer shelf and the slope was characterized by thePlethopeltid-Anomocarid Biofacies where Koldia, Kuljumbina, Plethopeltoides, Anomocariopsis, Paracoosia, Miranda, and Schoriecare weredominant, and agnostoid trilobites were less common. However, the Agnostid Macrobiofacies remained of primary importance in the trilobiteassemblages of the upper slope more removed from the margin of the carbonate platform. Lisogoragnostus, Proagnostus, Tomagnostella,Innitagnostus, Ammagnostus, Oidalagnostus, Acmarhachis, and Clavagnostus are characteristic of the assemblages of that macrobiofacies (Fig.12). At the beginning of Late Cambrian time, during the sea-level rise, Ceratopygidae became of major importance in addition to the agnostids in thetrilobite associations of the lower slope region. At that time the Agnostoid Macrobiofacies was dominated by Agnostidae (Innitagnostus,Proagnostus, Acmarhachis, and Glyptagnostus). Among polymerids of the Agnostid-Ceratopygid Biofacies, Proceratopyge, Aplotaspis,Schmalenseeia, Paradamesella, Acrocephalites, Acrocephalaspis, Eugonocare (Pseudeugonocare), and Onchonotellus are prominent (Fig. 13). In

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