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Ecological Succession' in Upper Jurassic Hardgrounds from Central Poland

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Ecological Succession' in Upper Jurassic Hardgrounds from Central Poland ACT A PAL A EON T 0 LOG IC A POLONICA Vol. 24 1979 No.4 MICHAl.. GRUSZCZy:flSKI ECOLOGICAL SUCCESSION' IN UPPER JURASSIC HARDGROUNDS FROM CENTRAL POLAND GRUSZCZyr<:lSKI, M.: Ecological succession in Upper Jurassic hardgrounds from Central Poland. Acta Palaeont. Polonlca, 24, 4, 42~~O, December 24, 1979. Analysis of ecological successions in Upper Jurassic hardgrounds at Skork6w (southwestern Holy Cross Mountains), central Poland, made it possible to test the general model of ecological succession proposed by Goldring and Kamler­ czak (1974). An ecological succession is not really evident in calcarenitlc hard­ grounds, except when developed on pelletal limestones. This is probably due to the high rate of cementation preventing community maturation. A succession is best developed in calcilutltic hardgrounds. Two additional criteria indicative of substrate consolidation are recognized: mode of oyster attachment and bi­ valve boring into filled formed crypts. A change from boring to nestling Is re­ corded in some bivalves. Influence of different hardground-microhabitats on the boring pelecypod associations is recognised. Key w 0 r d s: ecological Sliccession, hardgroun<is, Upper Jurassic, ea'rly ce­ mentation. Mtcha~ Gruszczyl\skt. Polska Akademta Nauk, Zaklad Paleobtolo17tt, 02-089 War­ 8zawa, al. Zwirkt i Wtgury' 93, Poland. Recettled: October 1978. INTRODUCTION The present paper is aimed to test and further refine the general model proposed by Goldring and Kazmierczak (1974) for ecological succession in hardgrounds. Kazmierczak and Pszczolkowski (1968) investigated Upper Oxfordian and Lower Kimmeridgian hardgrounds southwest of the Holy Cross Mts (for geological setting see Kutek 1968), recognized fossil "communities" associated with various types of hardgrounds. Some hardgrounds from that area were also studied by Kutek and Radwanski (1967) and Ronie­ wicz and Roniewicz (1968). Ecological succession and early cementation in Lower Kimmeridgian calcareous concretions from Mt. Policzko (W Holy Cross Mts), were studied by Kazmierczak (1974). Numerous hardgrounds occur in various types of Upper Jurassic Ii"': mestones of the southwest Holy Cross Mts. Well exposed hardgrounds . appear especially common in the Skorkow area (fig. 1). Twenty formed 430 ~CHA~ GRUSZCZy8SKI Fig. 1. Schematic map' of Upper Jurassic deposits from Skork6w vicinity. Kielce region. a large expOsures b other outcrops; J - Upper Jurassic (dotted), K- Cre­ taceous, Q- Quaternary. the basis for testing GoldriIllg and Kazmierczak's model, five of ~hich (figs 4 and 9) served as a basis for the reconstruction of organic associa­ tions inhabiting various types of hardgrounds. METHODS Over 150 polished slabs'cut normal to the hardground surface supplied the empirical data for the analysis of ecological succession and reconstruction of organic associations. Spatial structure of the associa­ tions was determined by the Schwerdtfeger's. (1968) method (With some ~ modifications). In every polished slab, a 2 cm thick interval including the hardground surface was diVided into 2 X 2 cm squares, and all components of the organic associations were counted. Slabs that did not allow unequivocal recognition of the spatial structure of the association as a random, cluster, or regular one, were removed from the sample set. The last generation of borings, distribution of the encrusting orga­ nisms, and intensity of the bioturbation were studied in every polished slab using the method introduced by Palmer and Palmer (1977) for Middle Ordovician hardground. ECOLOGICAL SUCCESSION AND LITHIFICATION Only a single investigated hardground (fig. 2) has developed in calci­ rudites, namely in chalky limestones composed of lumps, pellets, onkoids, ooids, and bivalve, gastropod, and brachiopod shells forming layers or lenses in places. Most investigated hardgrounds occur in calcarenites; the latter include various oolites (wackestones, packstones, grainstones), fine­ grained oolites, micro-oolites, and pelletal limestones. Finally, there are ECOLOGICAL SUCCESSION IN JURASSIC HARDGROUNDS 431 also some hardgrounds in calcilutites, represented by micritic limestones with micro-oolitic increments and shaly micritic limestones. In order to recognize ecological succession in -the hardgrounds exa­ mined, the relationship,' of organisms to substrate lithification (fig. 3) LI TH IFI CAT I ON plastiC;loose fir m I hard I undefinied burrows <OArenicolites t>:pe burrows ::=- ~.mpl. 00"""'=1 .c:=Thalassiroides type burrows ~ -=:: bivalve borings I. -=:: "Trypanites" borings I pedicolly attached brachiopods .I <:: oysters I serpulids Fig. 3. Organic structures and organisms associated with three stages of substrate Iithification was studied first. Early cementatio~ of the calcareous deposit leading ultimately to hardground formation may go either rapidly, or gradually and slowly (compare Goldring and Kazmierczak 1974). Three successive stages of substrate consolidation can be recognized irrespective of the rate of cementation. These are: 1. plastic or loose, 2. firm, 3, hard. In addition to the criteria given by Goldring and Kazmierczak (1974) for recognising substrate consolidation, two other criteria are proposed in the present study: 1) boring bivalves settling burrow openings (pI. 5: 4) and bored into older borings filled with shell detritus, indicating rapid lithification of the crypt fill (pI. 5: 5 and 6); 2) the mode of oyster attach­ ment. In the investigated hardgrounds, oysters attached to a firm sub­ strate with as large as possible area of their shells; the near-surface crust under tile shell is often fine-grained and reddish. If the substrate was hard, the oyster-shell attachment area is much smaller. Oyster shells attached to firm substrate are commonly bored by bivalves whereas those attached to hard substrates are usually not bored. This may reflect the best sites for settlement of boring bivalves i11 the case of firm substrates. ORGANISMS AND ORGANIC STRUCTURES ASSOCIATED WITH THE HARDGROUNDS Plastic or loose substrate: (1) Undefined burrows are usually poorly expressed. They range in diameter from some millimeters to a few centimeters; i 432 wnCHA~ GRUSZCZY~SKI (2) ThaZassinoides-type burrows include ~oth the Th. paradoxicus (Spongelimorpha paradoxical and Th. suevicus (S. suevica) morphotypes (cf. Fiirsich 1973; Kennedy 1967). Th. paradoxicus occurs in both plastic and firm substrates, while Th. suevicus appears restricted to firm substrate. / Firm substrate: (1) ThaZassinoides-type burrows; (2) Arenicolites-type burrows which include U-shaped channels rang­ ing in diameter from 0.6 to 1.5 cm. They never penetrate deeper than 5 cm from the hardground surface. These rare trace fossils are probably to be attributed to some polychaetes; (3) Simple burrows include three morphotypes: I- straight or slightly curved burrows (pI. 5: 1), distributed perpendicularly to the hardground surface. They range up to 2 cm in diameter and 8 cm in length; II­ short, bifurcate (reverse T-shaped) (pI. 5: 2) or geniculate (L-shaped) burrows extending some 2 cm below a hardground. Their diameter is approximate 0.5 cm at openings; III - single, Irregularly-shaped burrows usually with chisel-like distal ends (pI. 5: 3). They range up to 0.5 cm (opening diameter) and to 3 cm in length. Morphotype I resembles the burrows of some modern predatory crabs from the Bahamas (Shinn 1968) or Dca crabs (Basan and Frey 1977). (4) Bivalve borings were produced mostly by Lithophaga and Litho­ phaga-like pelecypods. Other boring pelecypods occur in minor amounts. Because of the lack of body fossils extracted from the rock, the Litho-. phaga-like pelecypods cannot be identified and hence, they are refered to as form "x·". Their borings are globose or bulgy in shape (their length to width ratio ranges from 0.9 to 1.1). All cross-sections of "x" and "x"-like borings are circular; . (5) "Trypanites" borings include three distinct morphotypes (d. 13ro­ mely 1972): I- straight or undulating tubes ranging from 0.1 to 0.3 cm in diameter and 6-8 cm in length (= "Trypanites" weisei) (pI. 6: 3; pI. 7: 1b); II - short (at most 1 cm in length), wedge-ended (pI. 8: 3), sometimes U-shaped channels approximate 0.2 cm in diameter. Similar borings are commonly regarded as analogous to those of the modern polychaetes PoZydora and PotamiZZa (d. Holder and Hollman 1969); III - "Trypanites" borings - a group close to the first one except that the former does always show wide, globose endings (pI. 8: 2b); (6) Pedically att;lched brachiopods are represented by ZeiZZeria hu­ meraZis, Epithyris subseHa, and SeptaZiphoria pinguis; (7) Oysters include representatives of Nanogyra {N. nana), Exogyra, and Liostrea. Their shells are commonly considerably damaged and hence, they.can hardly be identified. ECOLOGICAL SUCCESSION IN JURASSIC HARDGROUNDS 433 Hard substrate: (1) Bivalve borings; (2) "Trypanites" borings; (3) Pedically attached brachiopods; (4) Oysters; (5) Serpulids include three morphotypes: CycZoserpula (tube forms a circle), Tetraserpula (tube tetragonal in cross-section), and Dorsoserpula (tube round with a suture). ECOLOGICAL SUCCESSION IN CALCmUDITIC HARDGROUNDS Only one hardground of this type has been re,corded (fig. 2: 1). It is unique in that it has formed at a layer of calcareous nodules and asso­ ciated with shells of Diceras and Trichites. Close to and ~t those shells, there are polychaete borings represented mostly by straight- or U-shaped channels. Except for \Some singu~ar, U-shaped, (polychaete channels approximating 1 cm in
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