The main aim of this thematic issue is to present new high-resolution taxonomic data and ecologic interpretations that are currently available for the Kellwasser Crisis and other Late Devonian events from different parts of the Devonian world. Nine contributions present results of studies of eastern Laurussian epeiric successions in the framework of the international project “Ecosystem Aspects of Late Devonian Biotic Crisis

Baliński, Andrzej

Olempska, Ewa

Racki, Grzegorz

Acta Palaeontologica Polonica

English

Balinski, A

Olempska, E.

Racki, G.

Biblioteka Nauki - repozytorium artykuÅÃ³w

Biotic responses to the Late Devonian global events: Introductory remarks

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             Title: Biotic responses to the Late Devonian global events: Introductory remarks  Author: Andrzej Baliński, Ewa Olempska, Grzegorz Racki  Citation style: Baliński Andrzej, Olempska Ewa, Racki Grzegorz. (2002). Biotic responses to the Late Devonian global events: Introductory remarks. "Acta Palaeontologica Polonica" (2002, vol. 47, no. 2, p. 186-188).  Biotic responses to the Late Devonian global events:Introductory remarksANDRZEJ BALIŃSKI, EWA OLEMPSKA, and GRZEGORZ RACKIInvestigations of the Late Devonian successions have provideda great deal of evidence to support the hypothesis that thebiospheric turning point near the Frasnian–Famennian (F–F)boundary was one of the most significant in Phanerozoic time.Considerable attention and debate have been focused on deter−mining the cause(s) of these biocrises, and various mechanismshave been proposed for the extinction of a wide range of benthic,primarily low−latitude stromatoporoid−coral reefs, and pelagicorganisms. The widely recognized stepwise pattern of extinc−tion in major fossil groups in the late Frasnian argues against asingle catastrophic (“bedding−plane”) mass killings of biota atthe end of the Frasnian (see concept of a prolonged Kellwasser[KW] Crisis; Schindler 1993). However, many aspects of thismajor biotic turning point remain conjectural, including timingand magnitude of ecosystem changes and mediated extinctiondynamics. The overall biotic succession is far from reliably doc−umented. Among the better known pelagic faunas, a conodontnear−extinction and goniatite crisis are traditionally fixed pre−cisely at the F–F boundary.The main aim of this thematic issue is to present new high−resolution taxonomic data and ecologic interpretations that arecurrently available for the Kellwasser Crisis and other Late De−vonian events from different parts of the Devonian world (seelist of paper on the back cover of the issue). Nine contributionspresent results of studies of eastern Laurussian epeiric succes−sions in the framework of the international project “EcosystemAspects of Late Devonian Biotic Crisis”. This project was sup−ported by the State Committee for Scientific Research (KBNgrant no. 6 P04D 024 13 to G. Racki), and conducted jointly in1998–2000 by scientists from the Silesian University and the In−stitute of Paleobiology of the Polish Academy of Sciences, incooperation with specialists from Russia, Belgium, France, andUSA. In this issue, there are six complementary articles, includ−ing those by invited authors from USA, China and France. A re−lated paper to be soon published in this journal by Jerzy Dzikwill provide a comprehensive synthesis of the emergence andcollapse of Frasnian conodont and ammonoid communities inthe Holy Cross Mountains. Noteworthy is that this project is acomprehensive continuation of the previous KBN grant forG. Racki, dealing exclusively with the F–F brachiopod faunas,presented recently in the Devonian thematic issue of ActaPalaeontologica Polonica (Racki and Baliński 1998). Theremaining stratigraphic, litho− and palynofacies and geochemi−cal results of this project are presented in three articles in the si−multaneously appearing special issue of Palaeogeography,Palaeoclimatology, Palaeoecology (Racki and House 2002).The regional case studies in this volume refine and empha−size various aspects of global−scale biotic changes in Late Devo−nian marine settings, arranged from west to east (see Fig. 1).From western Laurussia, Day and Over described a unique low−est Famennian brachiopod survivor fauna from an offshore set−ting in the Appalachian Basin. The cyrtospiriferid−productoiddominated benthos was typical of the remainder of the earlyFamennian in many Laurussian basins (see Baliński). Feist’s in−vestigation of the trilobite fauna from the F–F boundary beds ofMorocco documents striking similarities in composition andevolutionary trends in contemporaneous faunas from North Af−rica and several European sections. This strongly argues againstwide oceanic separations of Laurussia and Gondwana postu−lated by workers citing published paleomagnetic data.The main group of eight articles document diversity of fau−nal records from different parts of the south Polish carbonateshelf of the south−eastern Laurussia, based mostly on data fromexposures in the Holy Cross Mountains and the Cracow area(Baliński, Ginter). Vishnevskaya et al. present the first detaileddocumentation of a world−wide bloom of siliceous radiolarian−sponge biota during the KW Crisis (see also Yudina et al.). Thelatters also discuss paleooceanographic and evolutionary impli−cations of the radiolarian−sponge bloom for both zooplanktonand benthos. Papers by Casier et al. and Olempska focus on re−cords of benthic and planktic ostracods. A combined, micro−facies−palaeontologic study of a fore−reef succession by Casierand his colleagues points to a regressive episode that startedclose to the fatal F–F stage boundary interval, paired with veryhigh rate of extinction in this well−oxygenated habitat. The pa−per by Olempska illustrates the role of entomozoaceanostracodes in tracing global climate and sea−level changes, andtheir effect on the ocean’s water masses during the Upper KWEvent. The entomozoaceans remained virtually absent during along time interval after the stepped end−Frasnian collapse, andreappeared late in the early Famennian as new species of refugialineages.Krawczyński describes Frasnian gastropod reef− and la−goon−dwelling faunas, and their close link with shallowing−upward cycles and sedimentary evolution of the carbonate com−plex. In a high−resolution study of the brachiopod faunas,Baliński demonstrates that the post F–F survival interval wascharacterized by low diversity and low frequency cyrto−Acta Palaeontol. Pol. 47 (2): 186–188, 2002Acta Palaeontologica Polonica 47 (2): 186–188, 2002Introductionspiriferid–rhynchonelloid or cyrtospiriferid–productoid assem−blages that feature stunted phenotypes characteristic of dis−turbed and stressed habitats during the survivor phase followingthe extinction. Preliminary results from studies of Givetian toFrasnian bryozoan faunas are discussed in biogeographic andglobal event contexts by Morozova et al.The paper of Głuchowski documents three successiveFamennian repopulation faunas characterizing post−extinctionrecovery of crinoids (defined with the aid of stem−based taxa),and shows a close correspondence between successive repopu−lation radiations and eustatic transgressive−regressive cycles.Ginter outlines the impact of the F–F biotic crisis on thechondrichthyan fauna. He suggests that global cooling is re−sponsible for the crash of the Frasnian subtropical extreme−stenothermal phoebodont sharks.Two other articles focus on contrasting F–F environmentalsettings in different parts of the East European Platform. Theevolutionary dynamics of successful rhynchonellid faunas af−fected by intermittent transgressions into the central region ofthe platform is analyzed by Sokiran, especially in the context oftheir early Famennian expansion. Newly refined stratigraphicevidence is available from a deep−shelf succession of the subpo−lar Urals in the study by Yudina et al. Many well known bioticand geochemical F–F phenomena (icriodontid and biosiliceousacmes, positive carbon isotopic shift) support the developmentof an unstable eutrophicated and oxygen−depleted crisis ecosys−tem in the tectonically disturbed Ural Ocean. The northernmostLaurussian sequence exhibits close similarities to the intra−shelfbasin succession in southern Holy Cross Mountains.The paper by Ma et al. presents interesting data concerningF–F extinction and recovery events among tetracorals, brachio−pods and ostracods in central Hunan, South China. Finally, re−view of literature−based data on distribution of Cyathaxonia cor−als by Wrzołek indicates that the Devonian was a time of rela−tively high diversity of this deeper marine fauna, significant de−crease in diversity of these corals this author sees rather at theGivetian–Frasnian boundary. The author proposes the new term“silent taxa” for those taxa, which are absent in significant inter−vals of their total range, exemplified by the Frasnian record ofthe Cyathaxonia coral group.Two general conclusions emerge from the regional contribu−tions, discussed above: The records of crinoids and radiolarians show minorchanges across the F–F extinction interval. In those groups,their main taxonomic turnovers occurred either later (e.g., atthe Devonian–Carboniferous boundary) or earlier (near theMiddle–Upper Devonian boundary). This is also seen in re−cords of some rugosan corals (Cyathaxonia faunas) andbryozoans. The records of sponges and gastropods remainsporadic. A biogeographic variability of extinction, survival and re−covery patterns, previously recognized in other major massextinctions (Erwin 1998), may be assumed at least for theF–F brachiopod and crinoid communities. Polish and NorthAmerican survivor and recovery brachiopod faunas are quitesimilar in terms of genera involved, but quite distinct fromthose reported from Russia and China. Webster et al. (1998)implied that the crinoids were relatively unaffected by theF–F crisis, although significant extinctions of crinoidsacross the F–F interval are clearly recorded in Holy Crossfaunas in Poland. In a taxonomic context, suggests that thepost−extinction renewal of the brachiopod fauna was quiterapid (ca. 1.5–2.0 Ma), but a significantly delayed reboundand rediversification is known among other faunas, e.g.echinoderms (Lane et al. 1997; Erwin 1998).As an integrative biostratigraphic synopsis, the data in thisissue constrain some of the extinction interpretations and uncer−tainties. In particular, new taxonomic summaries from three De−vonian continents (Fig. 1) yield more precise biostratigraphiccontrol needed to determine timing of events occurring withinLate Devonian global events. However, better understanding ofcomplex biotic and environmental feedbacks leads to severalnew unresolved questions, and to following key broadly−definedpaleobiologic matters for future interdisciplinary research: Major uncertainty is still evident because of common hia−tuses in the critical F–F interval in many subtropical andnear equatorial carbonate platform sequences. The quantita−tive approach to estimation of the true stratigraphic end−points (see confidence intervals on fossil ranges, and reviewin Marshall 1998) should be applied to distinguishing be−tween sudden and gradual biotic change. Records of manygroups affected by the F–F extinctions are difficult to assessstatistically because phylogenetic–taxonomic relationshipsin many fossil groups remain unclear. This ambiguity is evi−dent in the study by (in press) in conodont apparatus−basedtaxonomic terms, as well as for rugosan faunas by Wrzołek. The UKW crisis did not particularly affect the phyto−plankton (Streel et al. 2000), but a significant unbalancedexcess of bacterially controlled productivity is a marked fea−ture (Joachimski et al. 2001), resembling the disorganizedbiological pump after end−Cretaceous mass extinction.A major disturbance episode of the trophic web in the photiczone is clearly coupled with suppression of major pelagicconsumers (conodonts, ammonoids), augmented by Ginter’sdata on top predator (shark) selective demise. Sudden lat−eral shifts in prolific pelagic faunas (from radiolarian tohomoctenid−entomozoid to cephalopod communities) andcontribution of potentially tremendous diatom productivityto the F–F biosiliceous event are also puzzling (see Schieberet al. 2000). Investigation of survival communities and envi−ronments should yield a look into post−extinction strategiesand biogeochemical attributes. In a global ecosystem−geotectonic framework, reliable bi−otic and geochemical evidence is required to document andanalyze the timing of climatic changes (see discussion inStreel et al. 2000), especially for suspected Kellwasser cool−ing pulses, and a resolution of the nutrient sources drivingincreased primary biological productivity that induced geo−graphically variable anoxia (Yudina et al.) at different timesin the Late Devonian. Greatly refined studies of taxon ranges are required from thewidespread Middle Devonian carbonate platforms to theF–F carbonate crisis, particularly related to the many pulsesof transgression, anoxia and reef losses known over this pe−riod. McGhee (2001) recently suggested that a climatic keyto the KW Crisis enigma may be related to the Alamo impactin the overlooked early Frasnian timespan.http://www.paleo.pan.pl/acta/acta47/app47−186.pdfINTRODUCTION 187ReferencesDzik, J. (in press). Emergence and collapse of the Frasnian conodont andammonoid communities in the Holy Cross Mountains. Acta Palaeonto−logica Polonica.Erwin, D.H. 1998. The end and the beginning: recoveries from mass extinc−tions. Trends in Ecology & Evolution 13: 344–349.Golonka, J., Ross, M.I., and Scotese, C.R. 1994. Phanerozoic paleogeo−graphic and paleoclimatic modeling maps. In: A.F. Embry, B. Beau−champ, and D.J. Glass (eds.), Pangea; global environments and re−sources. Canadian Society of Petroleum Geologists, Memoir 17: 1–47.Joachimski, M.M., Ostertag−Henning, C., Pancost, R.D., Strauss, H., Free−man, K.H., Littke, R., Damsté, J.S., and Racki, G. 2001. Water columnanoxia, enhanced productivity and concomitant changes in  13C and34S across the Frasnian–Famennian boundary (Kowala—Holy CrossMountains/Poland). Chemical Geology 175: 109–131.Lane, N.G., Waters, J.A., and Maples, C.G. 1997. Echinoderm faunas of theHongguleleng Formation, Late Devonian (Famennian), Xinjiang−Uygur Autonomous Region, People’s Republic of China. Journal ofPaleontology 71 (Supplement to No. 2): 1–43.Marshall, C.R. 1998. Determining stratigraphic ranges. In: S.K. Donovanand C.R.C. Paul (eds.), The Adequacy of the Fossil Record, 23–53.J. Wiley & Sons, Chichester.McGhee, G.R. 2001. The “multiple impacts hypothesis” for mass extinc−tion: a comparison of the Late Devonian and the late Eocene. Palaeoge−ography, Palaeoclimatology, Palaeoecology 176: 47–58.Racki, G. and Baliński, A. (eds.) 1998. Brachiopods and the Frasnian–Famennian biotic crisis. Acta Palaeontologica Polonica 43: 135–411.Racki, G. and House, M.R. (eds.) . 2002. Late Devonian biotic crisis: eco−logical, depositional and geochemical records. Palaeogeography,Palaeoclimatology, Palaeoecology 181: 1–374.Schieber, J., Krinsley, D., and Riciputi, L. 2000. Diagenetic origin of quartzsilt in mudstones and implications for silica cycling. Nature 406:981–985.Schindler, E. 1993. Event−stratigraphic markers within the Kellwasser Crisisnear the Frasnian/Famennian boundary (Upper Devonian) in Germany.Palaeogeography, Palaeoclimatology, Palaeoecolology 104: 115–125.Streel M., Caputo M.V., Loboziak S., and Melo J.H.G. 2000. Late Frasnian –Famennian climates based on palynomorph analyses and the question ofthe Late Devonian glaciations. Earth−Science Reviews 52: 121–173.Webster, G.D., Lane, N.G., Maples, C.G., Waters, J.A., and Horowitz,A.B.S. 1998. Frasnian–Famennian extinction was a non−event for cri−noids, blastoids and bryozoans. Geological Society of America, Ab−stracts with Programs 30 (7): 30–31.AcknowledgementsWe would like to express our thanks to the following scientists whohave kindly assisted us by providing referee reports on manuscriptssubmitted for this Thematic Issue: Jonathan C. Aitchison, FrancoiseP. Bigey, Robert B. Blodgett, Denise Brice, Tomasz K. Baumiller,Jean−Georges Casier, Ivo Chlupač, Paul Copper, Jed Day, Jerzy Fedo−rowski, Richard A. Fortey, Jiri Frýda, Helga Groos−Uffenorde, Steve J.Hageman, Michael R. House, Gilbert Klapper, Krzysztof P. Krajewski,Norman M. Savage, James E. Sorauf, Jerzy Trammer, Gary D. Webster,and an anonymous reviewer.Paleogeographic models on p. 185 after Copper (1986), modified.Andrzej Baliński [balinski@twarda.pan.pl], Ewa Olempska [olempska@twarda.pan.pl], Instytut Paleobiologii PAN, ul. Twarda 51/55, PL−00−818 Warszawa, Poland;Grzegorz Racki [racki@us.edu.pl], Wydział Nauk o Ziemi, Uniwersytet Śląski, ul. Będzińska 60, PL−41−200 Sosnowiec, Poland.188 ACTA PALAEONTOLOGICA POLONICA 47 (2), 2002Fig. 1. Location of studied areas presented in this issue against the Late Devonian (363.0 Ma) palaeogeography (adapted from Golonka et al. 1994; courtesyof Jan Golonka).

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Biotic responses to the Late Devonian global events: Introductory remarks

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Biotic responses to the Late Devonian global events: Introductory remarks

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