Occurrence and paleoecological significance of lyssacinosid sponges in the Upper Cretaceous deposits of southern Poland
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Cretaceous lyssacinosid sponges (Hexactinellida) are rare and poorly recognized. This is the first description of lyssacinosid sponges from the Cretaceous of Poland. The sponges (including six species and three types of root tufts) come from the Upper Turonian–Lower Coniacian of the Opole Trough, Upper Campanian of the Miechów synclinorium, and Upper Campanian of the SE part of the border synclinorium. All localities lie southwards of the previous reports, widening thus the paleogeographic distribution of the group within the North European Province. Cretaceous lyssacinosids seem to be a useful tool in paleoecological interpretations. The presence of thin-walled lyssacinosids with root tufts indicates a soft substrate, slow rate of sedimentation, and calm and deeper water conditions.
KeywordsSponges Hexactinellida Lyssacinosida Paleoecology Upper Cretaceous Southern Poland
Sponges of the order Lyssacinosida Zittel, 1877 (subclass Hexasterophora Schulze, 1886) are a group of siliceous sponges of the class Hexactinellida Schmidt 1870, commonly referred to as “glass sponges”. Their oldest representatives are known from the Upper Proterozoic (Gehling and Rigby 1996). Mesozoic lyssacinosids are rare and poorly recognized. Few assemblages of these sponges are known from the Triassic of Utah and Nevada (Rigby and Gosney 1983), China (Wendt et al. 1989; Wu 1989) and Poland (Pisera and Bodzioch 1991; Bodzioch 1993, 1994). Jurassic lyssacinosids were described from Germany (Zittel 1877; Schrammen 1937), Morocco (Dresnay et al. 1978) and Argentina (Beresi 2003). Their presence in Triassic and Jurassic reef biota in the Alpine Region is documented by loose spicules (Mostler 1989; Delecat and Reitner 2005; Delecat et al. 2011).
Occasionally, lyssacinosids appear in Upper Cretaceous sponge assemblages of the North European Province, which consists mainly of other hexactinellids (Hexactinosida and Lychniscosida) and lithistids. Best known are Coniacian lyssacinosids from Bornholm (Mehl 1992; Brückner and Janussen 2005; Brückner 2006). However, single species of Lyssacinosida were also identified in the Cenomanian of Normandy (Moret 1926), Lower Cenomanian and Campanian of northwest Germany (Schrammen 1912; Salomon 1990), and from the Chalk of England (Bowerbank 1869; Reid 1968). Lyssacinosida have never been documented from the Upper Cretaceous of Poland.
Because the skeleton of lyssacinosid sponges is composed of unfused, or only locally connected spicules, their fossilization potential is definitely lower than that of other siliceous sponges with a rigid skeleton (e.g., Krautter 2002). The poor preservational state of these sponges in the Upper Cretaceous is the reason for their poor taxonomic recognition. Root tufts of lyssacinosid sponges composed of long basal spicules, occurring in the Upper Cretaceous deposits of southern Poland, were mentioned in the literature as sea grasses (Rutkowski 1965) or plants (Roemer 1870, pl. 30, fig. 2; Kędzierski 1995).
This paper aims to describe the lyssacinosid sponges from the Upper Turonian–Lower Coniacian of the Opole Trough, Upper Campanian of the Miechów synclinorium, and the SE part of the border synclinorium (Middle Vistula River section). In addition, their importance in paleoecological reconstructions is discussed.
Materials and methods
The studied specimens are hosted at the Institute of Geological Sciences of the Jagiellonian University, Krakow, collection no UJ220P, and in the Laboratory of Geology of the University of Lodz (collection no. ULXXIII).
In the extra-Carpathian Poland, Upper Cretaceous rocks crop out in the Southern Polish Uplands, in the Opole Trough, and in the Sudety Mountains (Fig. 1a). The so-called mid-Cretaceous eustatic transgression started in the middle Albian, covering rapidly most of its territory (Pożaryski 1960; Marcinowski 1974; Marcinowski and Radwański 1983, 1989). The initial facies variability of the Albian and Cenomanian was quickly followed by a uniform facies during the Early Turonian. With the exception of the Sudetes, where siliciclastic sedimentation prevailed, the rest of the area is characterized by limestone-marly facies (Central Polish Uplands); limestones are restricted to the Krakow Swell area, while in other regions opoka (siliceous limestone)/marly facies dominate (Marcinowski 1970, 1974; Walaszczyk 1992; Voigt et al. 2008).
The Cretaceous succession of the Opole Trough spans the Cenomanian to Middle Coniacian time interval (Walaszczyk 1992).
Rare lyssacinosid sponges occur in the Upper Marl Unit. All specimens are strongly pyritized with only single siliceous spicules preserved.
In the Miechów synclinorium, Cretaceous strata are represented by Upper Albian through to Lower Maastrichtian (Rutkowski 1965; Marcinowski 1974; Heller and Moryc 1984; Hakenberg 1986; Walaszczyk 1992; Świdrowska and Hakenberg 1999). The lyssacinosids were collected from three sections; two located in the southern part of the synclinorium (Rzeżuśnia and Strzeżów), and one in its northern region (Jędrzejów) (Fig. 1a, c).
The Rzeżuśnia section is an inactive quarry with exposed lower Upper Campanian [Inoceramus azerbaydajensis/Inoceramus vorhelmensis Interval Zone according to Jagt et al. (2004) and Walaszczyk et al. (2008)] opokas with cherts and marly intercalations. The relatively abundant fauna is dominated by inoceramids, gastropods, hexactinosid and lychniscosid sponges; less common are lithistid sponges and echinoids. Trace fossils are represented by Thalassinoides burrows. Lyssacinosids are very common, but only root tuft are preserved.
The Strzeżów section is an inactive quarry, with an Upper Campanian opoka succession, belonging to the “Inoceramus” inkermanensis and “Inoceramus” costaecus Zones (Jurkowska in prep.). The opokas are extremely fossiliferous, with abundant inoceramids, pectinids, gastropods, echinoids, and sponges (mainly hexactinosids and lychniscosids). Poorly preserved lyssacinosids appear throughout the succession.
The Jędrzejów section was a temporary road cut in the northern part of the city of Jędrzejów. The succession is composed of opokas and marls of the “Inoceramus” inkermanensis Zone to “Inoceramus” costaecus—“Inoceramus” redbirdensis Zone (Jurkowska, in prep.). The “I”. inkermanensis Zone is very fossiliferous, with abundant inoceramids, pectinids, belemnites, and echinoids. In the “I”. costaecus and “I”. redbirdensis Zones, echinoids and inoceramids become rare and big gastropods occur. Hexactinosid and lychniscosid sponges are numerous in the lower part of the succession but less common in the upper part where massive lithistids appear. Lyssacinosids are most numerous in the “I”. inkermanensis Zone, while in the “I”. redbirdensis Zone they are absent.
Microfacially, opokas from the studied sections represent packstone (Fig. 2b–e) with planktonic foraminifera and spicules of the siliceous sponges (mainly non-lithistid demosponges). In Rzeżuśnia, dispersed spicules of hexactinellid sponges also occur (Fig. 2c). In some thin-sections from Strzeżów, current alignment of spicules can be observed. Organic components also comprise fragments of bivalves and rare echinoids. An admixture of detritic material and glauconite was detected in Jędrzejów and Strzeżów; in Rzeżuśnia it is, however, insignificant.
All collected specimens of lyssacinosids are strongly limonitized. Rare, short fragments of siliceous or calcitized spicules have been found only in some specimens from Jędrzejów and Strzeżów. In all sections, root tufts type 1 are dominant. Large, fairly complete root tufts are preserved in life position while separated bundles of basal spicules lie horizontally.
SE part of the border synclinorium
In the SE part of the border synclinorium, the Upper Cretaceous succession is best exposed in the Middle Vistula River section (Pożaryski 1938; Marcinowski and Radwański 1983; Voigt et al. 2008) (Fig. 1a).
The Piotrawin section is an inactive quarry with a monotonous, poorly bedded succession of Upper Campanian opoka (Fig. 1c), which belongs to a local lithostratigraphic unit of Piotrawin Opoka (Walaszczyk 2004). Based on ammonites, Błaszkiewicz (1980) assigned the succession from this section to the Nostoceras pozaryskii Zone (=Nostoceras hyatti Zone according to Kennedy et al. 1992). Walaszczyk (2004) included its lower part into the “Inoceramus” altus Zone, whereas its middle and upper parts belong to the “Inoceramus” inkermanensis Zone.
The Piotrawin Opoka is extremely fossiliferous, with abundant siliceous sponges (Świerczewska-Gładysz 2006, 2012), ammonites (Błaszkiewicz 1980; Machalski 2012), belemnites (Kongiel 1962), inoceramids (Walaszczyk 2004) and non-ammonite molluscs (Abdel-Gawad 1986).
Microfacially, these opokas represent packstone (Fig. 2f) with foraminifera and spicules of siliceous sponges (mainly non-lithistid demosponges). Other bioclasts are fragments of bivalves, echinoderms, and rare bryozoans. Locally, in the upper part of the succession, current-aligned spicules are observed. Admixture of quartz and glauconitic grains is insignificant.
The lyssacinosid sponges are represented mainly by isolated root tufts. Bodily preserved sponges are much less numerous. Moreover, they are poorly preserved. All specimens are strongly limonitized and often only single spicules are visible. Fragments of siliceous spicules occur sporadically.
Lyssacinosid sponges from the studied sections
The studied material is dominated by isolated root tufts composed of basal spicules (spicules which protruded from the dermal surface and anchored sponges in the sediment).
Bodily preserved sponges are less numerous and, moreover, they are strongly deformed by compaction. They are usually incomplete, occasionally strongly fragmented with poorly visible single spicules. Better preserved specimens (described below) often possess only large spicules of the main (choanosomal) skeleton. Other taxonomically important spicules, such as autodermalia (megascleres supporting the dermal membrane directly), hypodermalia (megascleres lying under autodermalia), and supradermalia (megascleres protruding from the sponges surface), are rare. Microscleres, critical for the taxonomy of Recent taxa, have not been found. Due to their poor preservation, the taxonomic position of most of the sponges is uncertain.
Three specimens from Folwark.
The better preserved specimen is roundish, 35 mm in diameter (Fig. 3a). The round osculum on its top is 13 mm in diameter. The height of the sponge is unknown, as the specimen was compressed almost parallel to its axis. The dense choanosomal skeleton is composed of variably oriented diactines and diactine bundles, up to 6–7 mm long (Fig. 3b). Small, 1–2-mm-long, hypodermal diactines are arranged parallel to the wall surface (Fig. 3c). Single autodermal hexactines, supradermal orthotropal pentactines, paratropal pentactines (Fig. 3c), and fragments of few basal spicules occur (Fig. 3a).
Four specimens from Jędrzejów and 1 specimen from Strzeżów.
The sponges are probably cup-like (Fig. 3d) and more than 80 mm in height. All skeletons are strongly disintegrated. The spicules are scattered and their primary arrangement is occasionally unclear. The choanosomal skeleton is composed of isolated diactines (3–4 mm in length), diactine bundles (4–5 mm in length), and large hexactines (4–10 mm in size) (Fig. 3f). Supradermal pentacines are rare. Small autodermal hexactines (0.5–0.9 mm in size), found in a well-preserved fragment, form a dense and quite regular lattice (Fig. 3e). Numerous autodermal hexactines and few choanosomal hexactines (probably derived from the lower part of the body) were also observed between basal spicules. Straight or wavy basal spicules are thin, with an average thickness of ca. 0.1 mm. They comprise a dense, broad root tuft (Fig. 3d). Some of the basal pentactines are preserved with toothed anchors (Fig. 3g).
Three specimens from Strzeżów and 2 specimens from Jędrzejów.
The fragments are derived from thin-walled, probably cup-like sponges. The largest fragment, with a partly damaged wall, measures 160 mm. These sponges are characterized by an irregular choanosomal skeleton (Fig. 4a, b) and are composed mainly of giant hexactines, with thick rays (0.2–0.25 mm), and up to 16 mm long. The rays of some hexactines are curved (Fig. 4a). Variably orientated diactines, 0.1 mm thick and 4–8 mm long, are less common (Fig. 4b). Locally small autodermal hexactines, with rays up to 1 mm in length, are preserved (Fig. 4b).
Indeterminate genus and species A
Figure 4c, d
Five specimens from Jędrzejów and 1 specimen from Strzeżów.
The fragments are derived from thin-walled (ca. 1 mm thick) tube- and cup-like sponges, up to 50–90 mm in diameter and over 150 mm in height. Occasionally they are preserved with broken single basal spicules or small fragments of dense root tuft. The characteristic feature of these sponges is their choanosomal skeleton, composed of diactine bundles and isolated diactines, which run parallel and diagonal in growth direction (Fig. 4c). The diactine bundles are large, 20–30 mm long. Isolated diactines are smaller, 10–15 mm long on average. The hexactines, also occurring in the choanosomal skeleton, are 2–8 mm in size (Fig. 4d). Dermal spicules are absent.
Indeterminate genus and species B
Figure 4e, f
One specimen from Piotrawin.
This is a small, ca. 40-mm-high, thin-walled, cup-like sponge, with round to oval wall openings, 1–2 mm in diameter (Fig. 4e). The moderately dense skeletal lattice of this sponge is formed by long, thick (0.5–0.8 mm) diactine bundles, isolated diactines (0.1 mm thick and up to 3 mm in long) and rare hexactines, up to 2 mm in size (Fig. 4f). The diactine bundles run mostly in growth direction. Other spicules are not visible.
Indeterminate genus and species C
Three specimens from Folwark.
These are wall fragments (up to 3 cm in size), derived from cup-like sponges. The wall is thin (up to 1 mm) with irregularly distributed rounded or oval wall openings, 0.8–2 mm in diameter. The dense choanosomal skeleton consists of diactines, pointing in all directions, and of single hexactines. These spicules are poorly visible in a pyritized mass. The fragment of broad root tuft, preserved in the lower part of one of the specimen, is composed of thin (ca. 0.1 mm), straight spicules with a broken terminal part. Others spicules were not observed.
Root tufts type 1
- Figure 5a–d
Nine specimens from Folwark, 35 from Rzeżuśnia, three from Strzeżów, five from Jędrzejów, and four from Piotrawin.
The specimens are moderately dense, broad root tufts (Fig. 5a) composed of straight or slightly curved, thick (0.2–0.3 mm) spicules (Fig. 5d). The terminal parts of the spicules are not visible, but usually broken. The best-preserved specimens from Rzeżuśnia are very large, up to 250 mm long, and 160 mm wide (Fig. 5a). In the lower part of the root tufts, the spicules are loosely packed, the distance between them being up to 5–7 mm (Fig. 5b). A few specimens from Miechów synclinorium are represented by narrow (5–10 mm wide), separated bundles composed of few or several long spicules (Fig. 5c).
Root tufts type 2
Figure 5e, f
One specimen from Piotrawin and 2 specimens from Rzeżuśnia.
The specimens are composed of thin (0.05 mm thick), non-anchorate basal spicules which form narrow, separate, wavy tufts. The specimen from Piotrawin is composed of a fragment of the body and five separate tufts (up to 55 mm long and 2.5–3.5 mm wide) (Fig. 5f). The root tufts from Rzeżuśnia are broken and dispersed (Fig. 5e).
Root tufts type 3
One specimen from Folwark and two specimens from Piotrawin.
Root tufts of type 3 are stalk-like with nearly parallel straight spicules (diactines?), 0.1–0.15 mm thick (Fig. 5g). In the lower part of root tufts, the spicules are slightly dispersed. The best-preserved root tuft from Folwark is over 65 mm long, 5 mm wide in the upper part, and ca. 8 mm wide in the lower part. Specimens from Piotrawin are large, over 90 mm in length and 15 mm wide. Terminal parts of all specimens are broken.
Similar root tufts appear in several Recent lyssacinosids and in hyalonematid sponges (non-rigid hexactinellid sponges of the subclass Amphidiscophora Schulze, 1886). Sponges with stalk-like root tufts and some isolated stalk-like root tufts from the Upper Cretaceous of Germany were described as the sole fossil representatives of the genus Hyalonema Gray, 1835 (Mehl and Hauschke 1995). Basal spicules of living hyalonematid sponges usually have 4–8 teeth (Tabachnick and Menshenina 2002a). Due to the lack of the characteristic toothed anchors of basal spicules, the attribution of the specimen to Hyalonema is uncertain.
Most of the Recent Lyssacinosida live in the bathyal or even abyssal zone (e.g., Koltun 1967; Janussen et al. 2004; McClintock et al. 2005; Van Soest et al. 2007; Janussen and Reiswig 2009). Shallow-water (below 100 m) lyssacinosids are known exclusively from circumpolar seas (e.g., Koltun 1970; McClintock et al. 2005), the fjords of British Columbia (e.g., Leys et al. 2004), submarine caves in the western Mediterranean (Boury-Esnault and Vacelet 1994) and southern New Zealand (Leys et al. 2004). The occurrence of hexactinellid sponges in these places is possible only under special conditions such as: low temperatures, quiet waters, and low light levels (Leys et al. 2004; Whitney et al. 2005).
The distribution of ancient hexactinellids shows that they were more common in the neritic zone than their Recent counterparts. Development of hexactinellid sponges in the Late Cretaceous was correlated with high sea levels (Pisera 1999). It is also likely that the occurrence of lyssacinosids and other hexactinellids in Cretaceous European epicontinental seas, was related to the upwelling zones of the Tethys Ocean (see e.g., Mehl and Niebuhr 1995). The studied sections are located clearly southward of the formerly reported occurrences of lyssacinosids; the Tethyan influence could have been significantly higher.
The morphology and life function of lyssacinosids suggest that they were adapted to a low sedimentation rate and low turbulence, therefore they are located almost invariably in deeper shelf areas (e.g., Wendt et al. 1989; Pisera and Busquets 2002; Beresi 2003). Only some thick-walled sponges from the Triassic might have lived below the fair-weather wave-base but above the storm wave-base (Pisera and Bodzioch 1991; Bodzioch 1994). In all of the studied sections, lyssacinosids are represented by thin-walled forms, similar to deep-water forms known from the Coniacian of Bornholm (Brückner 2006). Recent thin-walled (<5 mm) lyssacinosid sponges occur at depth of ca. 100 m or deeper (up to 7,000 m) (Tabachnick 2002a, c).
Some modern lyssacinosid sponges live on a hard substrate, attached by a basal plate. Although basal plates occur in some fossil species (Pisera and Bodzioch 1991; Bodzioch 1994), they were not found in the studied sponges, suggesting their adaptation to soft-bottom conditions. Such adaptation is also suggested by the occurrence of lophophytal sponges (with root tufts) and numerous isolated long root tufts in the material studied (compare Brückner et al. 2003; Brückner 2006). The presence of lophophytal lyssacinosid sponges was used as an indicator of soft bottom conditions by Reid (1962) for the English Chalk Rock. Soft-bottom conditions in the studied successions is also suggested by the presence of rhizoidal lychniscosid and hexactinosid sponges (see also Tarkowski 1991; Świerczewska-Gładysz 2006), infaunal echinoids (Mączyńska 1968; Olszewska-Nejbert 2007), and some bivalves (Abdel-Gawad 1986).
The inferred life conditions of the studied sponges are also supported by other paleontological and sedimentological data. Packstone with foraminifera and spicules suggest calm water condition with periodic activity of weak currents. Weak currents favor the development of Recent hexactinellids (Leys et al. 2004; Whitney et al. 2005) and, consequently, are also inferred for the Cretaceous lyssacinosids.
Abundant lychniscosid and hexactinosid sponges, co-occurring with the studied lyssacinosids, indicate a deeper shelf environment (Świerczewska-Gładysz 2006). Moreover, some of these sponges [e.g., dish-shaped Ventriculites chonoides (Mantell, 1815) and Leiostracosia punctata (Schrammen, 1902)] indicate a very slow rate of sedimentation (Krautter 1997).
Thin-walled lyssacinosid sponges can only be preserved intact when quickly buried (Brückner et al. 2003; Brückner 2006). Most of the specimens studied herein have been partly destroyed due to a prolonged residence time on the sea floor, suggesting a slow rate of sedimentation in all of the successions. Quite often, only the root tufts that were buried in the sediment during the sponges’ lifetime have been preserved (Leys et al. 2007). The isolated root tufts of lyssacinosids and other non-rigid hexactinellid sponges were often described from other assemblages of fossil sponges, e.g., from Eocene of Catalonia (Pisera and Busquets 2002) or Permian of Texas (Rigby et al. 2007). The lack of well-preserved specimens and occurrence of dispersed spicules in the Campanian deposits result most likely from a slow rate of sedimentation and periodic weak bottom currents, which prevented the accumulation of the sediment and favored destruction of skeletons of dead sponges. Small specimens are more complete and may have been buried relatively quickly.
The significance of Lyssacinosida for bathymetric interpretation is confirmed by their distribution in the Cretaceous of the Opole Trough. Their occurrence in the Upper Marl Unit and absence from the underlying Marly Limestone Unit suggest deepening of the Opole Basin during the latest Turonian and Early Coniacian. This agrees well with the current bathymetric interpretation of the succession based on ichnofabrics (Kędzierski and Uchman 2001) and analysis of shark assemblages (Niedźwiedzki and Kalina 2003).
During the Campanian, the present-day Middle Vistula Valley area and Miechów area were situated in the Danish-Polish trough (Hakenberg and Świdrowska 1998). According to bathymetric interpretations based on non-cephalopoda molluscs from the Middle Vistula River section, Upper Campanian Piotrawin opokas were deposited in a mid to outer shelf setting (Abdel-Gawad 1986).
In the uppermost Campanian (“I”. costaecus—“I”. redbirdensis zones) of the Jędrzejów section (Miechów area) Lyssacinosida are not observed, suggesting shallower water conditions. This is confirmed by the replacement of hexactinellid sponges by lithistids in these zones (e.g., Reid 1968; Ulbrich 1974). In the stratigraphically equivalent deposits of the Middle Vistula River section, where lychniscosid and hexactinosid sponges are less common, Lyssacinosida have not been noted (Świerczewska-Gładysz, 2012).
Although lyssacinosids have been rarely reported from benthic assemblages of European Late Cretaceous epicontinental seas, it seems that they were more abundant than suggested by previous studies. So far, Late Cretaceous lyssacinosids have been recorded from in England, France, Germany, and Denmark. Their presence in the Upper Cretaceous deposits of southern Poland indicates that during the Late Cretaceous they had a wide paleogeographic distribution in the North European province.
In the studied collection Rossella rotunda Brückner, 2006, Rossella cf. bromlei Brückner and Janussen, 2005, and Chaunoplectella sp were recognized. The taxonomic position of three other species is impossible to determine, because the specimens are poorly preserved without diagnostic spicules. Due to the lack of rigid skeletons, most of the specimens are strongly disintegrated, and occasionally only root tufts, composed of thick, long spicules, are preserved. In the studied section, three types of root tufts were recognized.
Cretaceous Lyssacinosids can be used as paleoenvironmental tools. The presence of thin-walled forms with root tufts indicates a soft bottom, slow rate of sedimentation, and calm and comparatively deep water conditions.
We appreciate helpful comments by Dorte Janussen, Andrzej Pisera, and an anonymous reviewer. The authors are greatly indebted to Tadeusz Jurkowski, Jarosław Makieła, Krzysztof Paul, and Klaudiusz Salamon for help with the field work, to Beata Stożek for help with SEM photos, to Waldemar Obcowski and Grażyna Bartłomiejczyk for taking photographs of sponges, to Jarosław Szumny and Stanisław Olbrych for preparation of polished thin-sections, and to Aleksandra Wieczorek for linguistic corrections. The Institute of Geological Sciences Jagiellonian University Geology (grant No. PSP: K/DSC/000230) and Laboratory of Geology, University of Lodz (statutory fund, project 506/844) have supported this research.
This article is distributed under the terms of the Creative Commons Attribution License which permits any use, distribution, and reproduction in any medium, provided the original author(s) and the source are credited.
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