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(Radiolarians and Sponges) and the Late Devonian Biotic Crisis: the Polish Reference

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(Radiolarians and Sponges) and the Late Devonian Biotic Crisis: the Polish Reference Siliceous biota (radiolarians and sponges) and the Late Devonian biotic crisis: The Polish reference VALENTINA VISHNEVSKAYA, ANDRZEJ PISERA, and GRZEGORZ RACKI Vishnevskaya, V., Pisera, A., and Racki, G. 2002. Siliceous biota (radiolarians and sponges) and the Late Devonian biotic crisis: The Polish reference. Acta Palaeontologica Polonica 47 (2): 211–226. The radiolarian species Astroentactinia paronae, A. stellata, Trilonche echinata, T. grandis, T. nigra, Haplentactinia inaudita, and H. rhinophyuosa are common in late Frasnian to early Famennian rhythmic, calcareous−marly sequence of the southern Holy Cross Mts., Poland. They are known also from coeval abundant siliceous biota assemblages from the carbonate shelf of East European Platform including more than 150 taxa of radiolarians. However, in ecological terms, the moderately diverse Polish microfaunas (34 species of 12 genera) are more similar to these from Kolyma and Alaska, also marked by abundance of sphaerical entactiniids and near−absence of bilateral−symmetric Ceratoikiscidae and Palaeoscenididae. A succession of two distinctive siliceous sponges associations is established in the incipiently submerged Holy Cross carbonate platform: from an ephemeral, diverse, mostly rigid−skeletal lithistid−hexactinosan foreslope assemblage (initial phase of the late Frasnian Kellwasser Crisis), to long−lasting, basinal loose−skeletal hexa− ctinellid−demosponge faunas (appearing abundantly just prior the Frasnian–Famennian boundary in the late Palmatolepis linguiformis Zone). Such regional blooms of marine siliceous biotas, parallel to temporary retreat of calcareous biota, are demonstrated worldwide for the Kellwasser Crisis. These suggest probable causal links with cooling pulses and at least regional, volcanically induced eutrophication. Key words: Radiolaria, Porifera, palaeoecology, mass extinction, Frasnian, Famennian, Devonian, Poland. Valentina Vishnevskaya [[email protected]], Institute of the Lithosphere of Marginal Seas, Russian Academy of Sci− ences, Stromonetny per. 22, Moscow, 109180 Russia; Andrzej Pisera [[email protected]], Instytut Paleobiologii PAN, ul. Twarda 51/53, PL−02−089 Warszawa, Poland; Grzegorz Racki [[email protected]], Wydział Nauk o Ziemi, Uniwersytet Śląski, ul. Będzińska 60, PL−41−200 Sosnowiec, Poland. Introduction events (Schindler 1993), is used here. Earlier, Nazarov and Ormiston (1985) stressed the dissimilarity of comparative In the Late Devonian a carbonate shelf formed as a wide, evolutionary patterns between Palaeozoic radiolarians and the elongated belt on the pericratonic rim of the East European shelf−dwelling benthos. Similar conclusions were also pre− platform (Barskov 1994), stretching from the northern mar− sented by Kalvoda (1986), and Kiessling and Tragelehn gin of Europe (Pechora basin) to the southeast (throughout (1994), whilst very recent diversity compilations by Afana− Volga−Ural basin to Caspian−Dnieper−Donets basin), and sjeva (2000), Vishnevskaya and Kostyuchenko (2000), and then from the southern part of the Pripyat basin to the Polish Umeda (in press) present conflicting data. McGhee (1996) fragment (Holy Cross Mountains, Silesian−Cracow region). summarised dispersed evidence for an ecological shift toward The characteristic feature of these peripheral basins is the the dominance of siliceous biota, primarily within the benthos, presence of abundant siliceous microbiota (e.g., Nazarov during the F–F extinction interval. In fact, the southern part of 1988; Żakowa and Radlicz 1990; Vishnevskaya et al. 1993; the Holy Cross Mts. is distinguished by a locally abundant and Afanasjeva 2000). uniquely diverse late Frasnian siliceous sponge fauna (Rigby In the Holy Cross Mts., the first find of diverse Devonian et al. 2001), replaced by species−poor Famennian assem− polycystine radiolarians was made in marly Famennian strata blages, noted by Hurcewicz (1993). Thus, based on study of (see Racka 1984). In the present paper, the microfaunas from the reference Kowala succession (Fig. 1), the silicisponge as− the Frasnian–Famennian (F–F) passage at the Kowala section sociations are revisited herein in more details as well. Finally, are systematically described for the first time (see also Vish− a summary of the palaeoceanographic and evolutionary impli− nevskaya et al. 1997; Racki 1999). This led to a broader dis− cations is presented for both the F–F microzooplankton and cussion of this group response to the late Frasnian biotic turn− benthos (see Racki 1999). over, which is recorded primarily in collapse of stromato− V. Vishnevskaya and A. Pisera are responsible for radio− poroid−coral reef ecosystems. The general term “Kellwasser larian and sponge data, respectively, G. Racki for the strati− (KW) Crisis”, merging both the lower and upper anoxic KW graphic setting, other discussions are joint. Acta Palaeontol. Pol. 47 (2): 211–226, 2002 http://www.paleo.pan.pl/acta/acta47/app47−211.pdf 212 ACTA PALAEONTOLOGICA POLONICA 47 (2), 2002 A C L Early Pa. expansa D KOWALA Pa. trachytera K – Early bedded, ?TRANSGRESSION CARBONA Belorussian micritic ls. Pa. expansa High biohermal ls. Latest N J biostromal ls. Pa. marginifera icriodontids Ge F–F o coarse A sy n Holy Cross detrital ls. I Late BOUNDARY B c TE a li Mts fine-grained Pa. marginifera I s n in a detrital ls. l Early Pa. marginifera SHELF nodular ls. N Pa. rhomboidea homoctenids marly ls. SHALLOWING N marly shales Sub-Carpathian Pa. crepida E cherts Arch samples KM-1 studied H-4 M Manticoceras VOLCANISM high energy B events s Bodzentyn Late icriodontids appearance ss s ?SILICIC s Pa. triangularis FA ss PROGRESSIVE acme ssss s s UPLIFT disappearance ss REGRESSION ss Early–Middle ss marly sss Pa. triangularis ss H-3 see shales Kielce ss ss S³upia Nowa Pa. linguiformis Fig. 2 radiolarians marly lst. ss N micrites/ sssss s ssss Late cm sparstones s H-2 Pa. rhenana 80 Ky-3 A coquinas m cherts ?EarlyPa. rhenana I 60 crinoid debris KOWALA H-1 Pa. hassi 40 –Pa. jamiae cephalopods N G Pa. hassi 40 F icriodontids Permian S 0 E Pa. punctata radiolarians 010km entomozoids sponges Carboniferous A 20 D C articulates homoctenids B Amphipora Devonian R ? Pa. transitans UPPER KW A TRANSGRESSION articulates sssss sss ss 0 & ANOXIC EVENT Older Paleozoic £ysogóryss overthrust F Amphipora entomozoids Fig. 1. A, B. Location of the Kowala basin succession in Poland (A) and Holy Cross Mountains (B; see details in Racki 1993). C. Composite lithological section of the Late Devonian strata at Kowala (modified after Szulczewski 1996: fig. 8), the reference succession showing environmental evolution typical of intermittently drowned shelf from reef (units A–C) to slope (D–G) to intrashelf basin (H–L). Three sampled radiolarian−sponge intervals are arrowed. D. Sedimentary record of eustatic, anoxic and tectono−volcanic events around the Frasnian–Famennian boundary, with emphasis on selected biotic markers. Note blooms of siliceous biota, thought as promoted mostly by submarine exhalative volcanism paired with tectonically−driven shallowing, and a cooling pulse (see Racki 1999; Racki et al. 2002). Kowala basinal succession and cession (set H), four units are distinguished in the F–F strata (details in Racki et al. 2002; Figs. 1C, D and 2). There are: siliceous faunas H−1.—Variably−bedded, fossiliferous marly micrites are inter−stratified with graded detrital limestones (with broken Distinctive siliceous faunas have been found in the well− reef−builders and brachiopod shelly partings), and/or thick studied succession (e.g., Szulczewski, 1996; Joachimski et al. (up to 1 m) slump layers. 2001; Racki et al. 2002) at active Kowala quarry (Holy Cross H−2.—Thinly bedded marly limestones with shale interbeds Mountains, Central Poland; Fig. 1A, B). The Kowala section is comprise frequent, irregular, subnodular intercalations, up to a typical representative of the Late Devonian infilling of the 45 cm thick. Sparse fossil assemblages include both planktic southern (Chęciny−Zbrza) intra−shelf basin, adjacent to the cen− and benthic (mostly brachiopod) elements. Among the latter, tral palaeotopographic high (the Dyminy Reef). Within the diverse silicisponges are particularly characteristic of the over 150 m thick, dark to black rhythmic calcareous−marly suc− lower part of this unit (see Rigby et al. 2001). Fig. 2. Micropalaeontological characteristics of the upper Frasnian and Frasnian–Famennian passage at Kowala, based on the eastern quarry wall section ý from 1991 (section Kw). The material obtained from marly−shaly samples, supplemented by few conodont samples and thin sections for the unit H−3. Pro− ductive radiolarian (R) and spicule (S) samples studied (see Fig. 3 and Table 2) are arrowed. 1, tentaculitoids; 2, entomozoids; 3, large−sized palynomorphs; 4, goniatites; 5, ichtyolithes; 6, palmatolepid conodonts; 7, polygnathid conodonts; 8, ramose conodont elements; 9, phyllocarid remains; 10, benthic ostracods; 11, nanicellid foraminiferans; 12, sponge spicules; 13, rhynchonellid brachiopods ;14, chonetid brachiopods; 15, lingulids; 16, phosphatized bur− row fills and pellets; 17, agglutinated foraminiferans; 18, icriodontid conodonts; 19, echinoderm remains; 20, ramose fossils (a, amphiporid stromato− poroids; b, others [fragmented ?auloporids]); 21, sphaerical microfossils (?mostly calcispheroids). VISHNEVSKAYA—LATE DEVONIAN SILICEOUS BIOTA 213 22 157 154 H-3 FAMENNIAN (b) 148 20 HORIZON FRASNIAN (b) 144 Ass. BIOSILICEOUS set 141 140 ?MIXED Biofacies 137 Pa. linguiformis 136 B Ass. 18 136 A 135 TIONAL ? 133 TRANSI-
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