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Boring and Nestling Organisms from Upper Jurassic Coral Colonies from Northern Poland

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Boring and Nestling Organisms from Upper Jurassic Coral Colonies from Northern Poland Acta Palaeonto1,ogica Polonica Vol. 32 No. 1-2 pp. 83-104; pls. 33-40 Warszawa, 1987 ANDRZEJ PISERA BORING AND NESTLING ORGANISMS FROM UPPER JURASSIC CORAL COLONIES FROM NORTHERN POLAND PISERA A.: Boring and nestling organisms from Upper Jurassic coral colonies from northern Poland. Acta Palaeont. Polonica, 32, 1-2, 83+lW, 1967. The Upper Jurassic coral colonies collected from glacial deposits in northern Poland reveal abundant boring and nestling organisms. The most common borers were bivalves, among which 4 gastrochaenid species (Gasttochaena s. s. and Spenglerla) and 2 lithophagid species are described. Other borers included sipun- culid worms and sponges (7). The nestling fauna consists of the bivalves Hfatella (probably also a facultative borer), Pltcatula, and oysters, as well as serpulids. The assemblage of borers is suprisingly similar to Recent assemblages from coral reefs. The excellent preservation oi aragonitic bivalve structures allowed for a detailed examination of the internal structure of shells and boreholes. Basing on these features, new diagnostic characters are here proposed and a partial revision is undertaken. K e y w 0 r d s: Boring organisms, bivalves, taxonomy, Upper Jurassic, Poland. Andrzej Pisera, Zaktad Pakobfologft, Polska Akademfa Nauk, Al. twfrkf t Wtgu- q 83, 02-089 Warszawa, Poland. INTRODUCTION The coral colonies of Oxfordian-Kimmeridgian age (see Roniewicz 1984) gathered from glacial deposits of northern Poland show the presence of rich and diversified borings. The majority were executed by bivalves but other taxonomic groups were also responsible for some borings. In many cases, the boreholes contain their original inhabitants andlor se- condary dwellers. Boring and nestling bivalves of Jurassic age are known since long ago (see Phillips 1829, Eudes-Deslongchamps 1838, Buvignier 1843, 1852, Lori01 1888, 1892, 1893, Boehm 1893, Borissjak 1906) and numerous species have been described. The majority, however, are poorly, if at all, illustrat- ed and only inadequately described. Often, they are preserved as moulds or impressions only, which do not allow for a detailed examination of the shell. Moreover, they are oversplitted at the species level; Rollier (1914), for example, listed 47 specific names of the Jurassic lithophagid bivalves. 84 ANDRZEJ PISERA The excellent preservation of the currently investigated material permits, for the first time, to examine the interior of the shell and to classify these organism according to more recent standards. Carter (1978) and Freneix and Roman (1979) dealt recently with diffe- rent aspects of gastrochaenid taxonomy, evolution, and ecology. They examinted, however, mostly a Recent material. In the present paper I am trying to relate the results of these extremely important papers to an Upper Jurassic material.') The investigated material is housed in the Institute of Paleobiology, Polish Academy of Sciences, Warsaw, (ZPAL). Acknowledgements.-I am grateful to Dr. E. Roniewicz and Dr. J. Dzik (Institute of Paleobiology, Warsaw) for putting at my disposal the material for investigations. Dr. S. R. A. Kelly (Department of Geology, Cambridge University) supplied me with molds of the specimens housed in Cambridge, and Miss B. Pyrah (Yorkshire Museum, York) sent me the photograph of another specimen from the Museum collection. The staff of the VSEGEI (Leningrad) museum allowed me to investigate Borissjak's collection INVESTIGATED MATERIAL The original collection of coral colonies consists of 1023 specimens, most of them however are only small fragments. Nearly 26O/o of specimens show traces of different boring organisms; over 24O/o were bored by bivalves. Epifauna occurs on only 2O/o of specimens. In fact, the percentage of originally bored colonies had to be much higher, for the collection contains mostly fragments of larger colonies. More than 100 bivalves (boring and nestling) have been found and make up the basis for the present study. The collection consists mostly of Thamnasteria concinna; much less common is Actinaraeopsis exilis Roniewicz, while other species are repre- sented each by a single or few specimens (Roniewicz 1984). The richest collection of corals was gathered in the gravel pit at Ostromice (Wuster- mitz or Wusterwitz of German authors: Roniewicz 1984). Both the corals and the bivalves are exceptionally well preserved, in form of the original aragonite. AGE OF BORINGS As all the material comes from glacial deposits, there is no direct indication of the age of borings in coral colonies. Indirect evidence, howe- ver, points to the same age of corals and borings. In some natural cavities, 1) Note added in proof: The results of the important paper by Kleemann (1983) were not discussed here, as I was not aware that it was already published. BORING AND NESTLING ORGANISMS FROM JURASSIC CORALS 8 5 for example, ferruginous sandy to sandy carbonate rock occurs between coral colonies (Roniewicz 1984), and the same type of sediment fills up some of the borings which still contain intact bivalve shells; this indicates synsedimentary origin of the borings. Undamaged apertures of bivalve boreholes are preserved only on the original, and not on eroded, coral surfaces. If the corals were drilled only after deposition and redeposition, in a secondary environment, one could expect preservation of the original apertures also on the eroded coral surfaces, which is not the case. This observation also points to the same age of the corals and the borings. The most important evidence supporting such hypothesis, however, comes from the rare cases when a borehole, still containing its primary inhabi- tant, is completely overgrown by coral (pl. 39: 3) without any traces of disruption in the coral growth. More precisely, these cases indicate that the bivalves bored coral colonies still alive. BIVALVE BORERS In the investigated material, the majority of borings were made by bivalves, among which the most common were gastrochaenids (pl. 33: 3, 5-6) accompanied also by the genus Lithophaga. For a detailed descrip- tion of these bivalves see the systematic part of this paper. NON-BIVALVE BORERS AND BIVALVE SQUATTERS The most common non-bivalve borings are very small, tubular in shape (one to two millimetres in cross section), very irregular and interconnected in a chainlike pattern, sometimes bifurcated. They penetrate shallow below the surface and have only a single small, rounded opening (pl. 33: 1-4). They are similar to the Cretaceous trace fossil Entobia as well as to the Recent excavations of boring sponges. They are preserved on many investigated coral colonies but mainly on somewhat eroded surfaces, which does not allow to recognise the taxonomic position of the animal respons- ible for their origin. Other common boreholes are much larger, up to 5 centimetres long and up to 3 mm in diameter. They usually are gently curved and circular in cross section, and may penetrate deep into the coral substrate (pl. 33: 1-2, 5). Similar borings are produced in Recent corals by sipunculid worms (Warme 1975, 1977, MacGeachy and Stearn 1976). Basing on this morphological similarity and the ecological context, I am attributing these Jurassic borings to sipunculid worms. Many boreholes as well as some natural coral growth cavities in my material were inhabited by secondary dwellers (squatters). The most comm6n among them was the bivalve Hiatella (Pseudosaxicava) phaseolus 86 ANDRZEJ PISERA (Deslongchamps), which is described in detail in the systematic part. Where this species is found in a gastrochaenid or lithophagid borehole which still contains its primary inhabitant, this suggests that Hiatella was a squatter rather than borer. In some cases, however, it occurs alone in a borehole (pl. 37: 5) which differs from the gastrochaenid and litho- phagid borings; this suggests that the Upper Jurassic hiatellids already had some boring ability, at least the ability to reshape a primary borehole. Sometimes, three generations of inhabitants of the same borehole existed; a gastrochaenid borehole (with the shell inside) was colonized by Hiatella, which was finally succeeded by a young Lithophaga. Other common inhabitants of empty boreholes, as well as growth cavities of coral colonies, were serpulids (probably Glomerula gordialis (Schlotheim) (pl. 34: 5-4) and small spinose Plicatula sp. (resembling Plicatula spinosa Sowerby) (pl. 34: 1-2, 4). A few oysters colonized large natural coral growth cavities and mimicked their shape (pl. 34: 3). ECOLOGICAL REMARKS Roniewicz (1984) considered the lithology of the matrix and the growth forms of the corals, and postulated a low energy environment with high influx of detrital material from crystaline rocks as the original setting for these corals. According to her, the bottom was unstable sand or sandy mud with admixture of biogenic detritus where colonies were overturned only during periods of rougher conditions. In some cases, borings occur on all sides of the investigated colonies; generally, however, the lower and lateral surfaces were infested thus suggesting that most borers attacked overturned and already dead corals. Where a whole borehole, still containing its inhabitant, is entirely over- grown by the coral without any trace of disruption in the structure of the colony, it suggests that a bivalve (Gastrochaena (G.) cf. deshayesea) infested and lived within the growing portion of the coral. The assemblage of boring organisms from this Upper Jurassic material is surprisingly similar to the associations known from Recent shallow water
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