BULLETIN DE L'INSTITUT ROYAL DES SCIENCES NATURELLES DE BELGIQUE SCIENCES DE LA TERRE, 74: 119-127, 2004 BULLETIN VAN HET KONINKLIJK BELGISCH INSTITUUT VOOR NATUURWETENSCHAPPEN AARDWETENSCHAPPEN, 74: 119-127, 2004
Taphonomic and ethologie aspects of the ichnology of the Maastrichtian of the type area (Upper Cretaceous, The Netherlands and Belgium)
by Stephen K. DONOVAN & John W.M. JAGT
D o n o v a n , S.K.. & J a g t , J.W.M., 2004. - Taphonomic and ethologie au test en ayant d ’abondantes épines de l’échinide hôte enfouies dans ses aspects of the ichnology of the Maastrichtian of the type area (Upper tissus mous. L’organisme foreur Talpina cf. ramosa von Hagenow, Cretaceous, The Netherlands and Belgium).Bulletin de I 'Institut royal 1840 généralement conservé sous forme de moulages naturels dans des des Sciences naturelles de Belgique, Sciences de la 74: Terre, 119-127, coquilles décalcifiées, se présente sous forme de cavités dans l’huître 2 figs., 1 pi., Bruxelles-Brussel, March 31, 2004. - ISSN 0374-6291. Agerostrea ungulata (von Schlotheim , 1813). Trypanites cf. solitarius M ägdefrau, 1937 présente une morpholo gie inhabituellement aplatie dans laquelle la forme du forage a été en partie déterminée par la distribution des niveaux organiques dans la Abstract coquille de l’huître Rastellum macropterum sensu STENZEL, 1971 (Membre de Nekum, Formation de Maastricht). Three rare ichnotaxa from the type area of the Maastrichtian Stage (Upper Cretaceous) are described, and their morphology and palaeoecological Mots-clefs: Terriers, forages, “ enfouissement” , Maastrichtien, Pays- significance discussed; a fourth ichnotaxon shows an unusual mode of Bas, Belgique. preservation. The infill of an articulated lithophagid bivalve (Meerssen Member, Maastricht Formation) preserves the distinctive firmground burrow Arachnostega gastrochaenae B e r t l i n g , 1992. This is the first report of this trace fossil, probably generated by polychaetes, from the Introduction Cretaceous of northern Europe. The producer may have been grazing the inside of the bivalve shell while burrowing through its sediment infill. An Trace fossils are an understudied, yet diverse and significant indeterminate patellid limpet exhibits an additional example. Tubercles are recognised for the first time within the pits ofOichnus excavatus element of the biota of the Maastrichtian (Upper Creta Donovan & Jagt, 2002b (Meerssen Member, Maastricht Formation), ceous) in its type area in Liège-Limburg (Belgium) and indicating they were probably the result of embedment rather than boring. southern Limburg (The Netherlands)(D o r t a n g s , 1998; The producing organism may have obtained a firmer attachment to the test by having abundant spines of the host echinoid embedded in its soft D o n o v a n & J a g t , 2002a; J a g t , 2003) (Fig. 1). The present tissues. The boring Talpina cf. ramosa von Hagenow, 1840, commonly authors are attempting to correct this omission by describ preserved as natural casts in decalcified shells, occurs as holes in the ing previously unreported but significant trace fossils from oyster Agerostrea ungulata (von Schlotheim, 1813). Trypanites cf. the various members included in the Maastricht Formation. solitarius M ägdefrau, 1937, shows an unusually flattened morphology, in which the boring form has been determined in part by the distribution Herein, we describe three unusual and/or rare and morpho of organic layers within the shell of the oyster Rastellum macropterum logically distinct ichnofossils from this unit, each of which sertsu S t e n z e l , 1971 (Nekum Member, Maastricht Formation). shows unusual features related to preservation, plus add Key words: Burrows, borings, embedments, Maastrichtian, The Nether further observations to the original description of an ichno lands, Belgium. taxon that suggests a most peculiar behaviour. Terminology of the morphology of trace fossils follows H ä n t z s c h e l (1975). The philosophy of open nomencla Résumé ture follows B e n g t s o n (1988). Annotations within syno nymy lists follow M a t t h e w s (1973). All specimens are Trois ichnotaxa rares provenant de la région type de l'étage Maastrich- tien (Crétacé Supérieur) sont décrits; leur morphologie et leur significa deposited in the Natuurhistorisch Museum Maastricht, The tion paléoécologique sont discutées. Un quatrième ichnotaxon montre un Netherlands (NHMM); JJ = J.W.M. Jagt collection, LN mode de préservation inhabituel. Le remplissage d’un bivalve lithophage = L. Nelissen collection, MK = W.M. Felder collection. articulé (Membre de Meerssen, Formation de Maastricht) a conservé le terrier semi-consolidé caractéristique Arachnostega gastrochaenae BERTLING, 1992. C’est la première fois que cette trace fossile laissée probablement par les polychaetes, est signalée dans le Crétacé de l’Eu Systematic ichnology rope du Nord. L’animal à l’origine de ce terrier peut avoir raclé l’inté rieur de la coquille du bivalve pendant qu’il creusait dans le sédiment de remplissage. Une patelle appartenant à un patellidé indéterminable en IchnogenusArachnostega B e r t l in g , 1992 montre un autre exemple. Des tubercules ont été observés pour la première fois dans les cavitésd'Oichnus excavatus Donovan & Jagt, T y p e ichnospecies 2002b (Membre de Meerssen, Formation de Maastricht) indiquant qu’elles résultent probablement d’un “ enfouissement” plutôt que d’un Arachnostega gastrochaenae B e r t l in g , 1992, p. 179, by creusement. L’organisme responsable peut avoir été plus fermement fixé original designation. The only nominal ichnospecies. 120 Stephen DONOVAN & John JAGT
1994 Arachnostega gastrochaenae Bertling - F ü rsich et al., pp. 146, 161, pi. 3, figs. 1, 2, 4. ? 2001 Trac/¡«ewíega-Wilson & Taylor , p. 26, pi. 2, fig. 1. k m
M a t e r ia l Two specimens. An incomplete internal mould of an indeterminate lithophagid bivalve, NHMM JJ 12354, with A. gastrochaenae apparent within the moulds of both valves (PI. 1, Figs. 1, 2). The internal mould of the bivalve, preserved in a medium- to coarse-grained bio- calcarenite, is broken both anteriorly and posteriorly, preserving little detail of the internal surface of the mol lusc shell except for some coarse growth lines. The second specimen, NHMM MK 367, is an internal mould of an indeterminate patellid limpet of the type described by K a u n h o w e n (1898, p. 15, pi. 1, figs. 3-4), with A. gastrochaenae weakly developed (PI. 1, Fig. 3). Fig. 1 — Outline map of study area (redrawn and simplified after Jagt , 1999, fig. 1), showing political boundaries L o c a l it y a n d h o r iz o n (dashed lines), rivers and canals (solid lines) and the NHMM JJ 12354 is from ENCl-Maastricht BV quarry, city of Maastricht (M). Key to localities: 1 = ENCI- south of Maastricht, southern Limburg (The Nether Maastricht BV quarry; 2 = Ankerpoort-’t Rooth quar lands), base of subunit IVf-4, Meerssen Member, Maas ry; 3 = temporary sections in Albertkanaal; 4 = CBR- tricht Formation. Upper Cretaceous; uppermost Maas Romontbos quarry. The inset map of the Netherlands (Nl), Belgium (B) and Germany (D) shows the ap trichtian (for simple lithostratigraphic section, seeJ a g t proximate position of the main map (box). et a l, 1998, figs. 1, 2). The label of NHMM MK 367 states “Nekami, Bemelen [now known as Ankerpoort- ’t Rooth quarry], ?Nekum Member,” Maastricht Forma tion. Upper Cretaceous; upper Maastrichtian (Fig. 1).
D ia g n o s is (From B e r t l in g , 1992, p. 179.) “ Irregular elongate and D ia g n o s is net-like burrows in sediment fills of shells. Visible on the (After B e r t l in g , 1992, p. 180). “ Ramified burrows on the surface of internal moulds. The size of the meshwork may surface of internal moulds with an oval cross-section, vary from microns to centimetres, depending on the shell- which increases slowly in diameter by a factor of 5 to 10, bearing and the burrow biota involved.” approximately. At ramifications, the bent main burrow is not reduced in size. Lateral burrows mostly have a smaller
R e m a r k s diameter. Polygonal areas on the surface of internal moulds Arachnostega is a distinctive network of channels that is may be produced by the unification of ramified burrows.” superficially reminiscent of structures such as the clionid sponge boring Entobia B r o n n , 1838, although lacking D e s c r ip t io n swollen chambers and not penetrating a calcareous Based mainly on NHMM JJ 12354. Burrows more inten shell or limestone substrate. That Arachnostega itself is sely developed adjacent to the right(?) valve (PI. 1, Figs. 1, not a boring was only determined by the astuteness of 2) than leftf?). Burrows best seen on surface of moulds, B e r t l in g (1992, pp. 182-183), who recognised that mor preserved in a surface layer that has, in part, peeled away phologically identical burrows occurred in unusual firm- on NHMM JJ 12354 (PI. 1, Fig. 2), revealing less common ground settings within sediment-filled shells of the in- burrows below. Burrows of incomplete circular or ellipti faunal bivalve Mya sp. within the North Sea Basin, cal section, preserved as grooves with overhanging edges generated by more than one taxon of polychaete annelid and not infilled, individual burrows varying fromc. 0.1 to (R e in e c k , 1980). However, Arachnostega has only been 0.6 mm in maximum diameter; broader sections of bur formally recorded hitherto from the fossil record of the rowsi?) maY be a taphonomic artifact. Larger diameter Jurassic of northern Europe (B e r t l in g , 1992; F ü r s ic h et burrows forming reticulate pattern at surface of mould. al., 1994) and the Cretaceous of the Middle East(W il s o n & T a y l o r , 2001). R e m a r k s This is the first reported occurrence of this distinctive, but
Arachnostega gastrochaenae B e r t l in g , 1992 cryptic, ichnotaxon from the Maastrichtian of the type (PI. 1, Figs. 1-3) area. W il s o n & T a y l o r (2001, p. 26) recorded it from approximately coeval sedimentary rocks of the Qahlah * 1992 Arachnostega gastrochaenae Bertling, pp. 177, Formation in Oman, uniquely preserved in mudstones in 179-185, figs. 2, 3. filling Gastrochaenolites isp. borings. However, the unu 1993 Arachnostega - T odd, p.417. sual mode of occurrence ofA. gastrochaenae within in Ichnology of the type Maastrichtian 121
filled shells and borings prevents its identification except cimens considered herein include the holotype pit in within mouldic material, so it is probably more widespread NF1MM MK 4689 (PI. 1, Fig. 6, arrowed), and selected in the Mesozoic and Cainozoic than has been reported paratypes, including 17 other pits in NHMM MK 4689 hitherto (see also comments inF ü r s ic h et al., 1994). (PI. 1, Figs. 4, 6, 7) and all pits in NHMM JJ 699 (at least Although present internally, the best development of 61 individual pits). burrows is undoubtedly on the surface of the internal mould, where movement would have been constrained by L o c a l it y a n d h o r iz o n the molluscan shell. This may indicate that the producing Upper 5-10 m of the Meerssen Member, Maastricht For polychaetes(?) were in some way grazing the interior of the mation, formerly exposed along the Albertkanaal at shell. They may have been feeding on non-photosynthetic Vroenhoven-Riemst, Limburg (Belgium) (Donovan & microbes that had been living on the inside of the shell. J a g t , 2002b, fig. 1). Upper Cretaceous; uppermost Maas trichtian (Fig. 1). IchnogenusOichnus B r o m l e y , 1981 D ia g n o s is T y p e ichnospecies (Slightly revised after Donovan & Jagt, 2002b, p. 69.) Oichnus simplex B r o m l e y , 1981, p. 60, by original des Circular to elliptical, non-penetrative Oichnus, almost ig n atio n . invariably with a broad, high, raised central boss. Aper ture of pit overhanging and walls concave. O t h e r ichnospecies Oichnus asperus N ielsen & Nielsen,2001 ; O. coronatus D e s c r ip t io n Nielsen & Nielsen, 2 0 0 1 ; O. excavatus D o n o v a n & (Further to features described in Donovan & Jagt, J a g t , 2 0 0 2 b ; O. gradatus Nielsen & Nielsen, 2 0 0 1 ; 2002b.) All surfaces of pits bearing sculpture of irregu O. ovalis B r o m le y , 1993; O. paraboloides B r o m le y , larly distributed to well-ordered tubercles, commonly 1981. very small (granular), although may be present in two sizes (PI. 1, figs 4, 6, 7). In at least one shallow pit, small D ia g n o s is tubercles are arrayed around larger tubercles in a circle. (After D o n o v a n & P ic k e r ill , 2 0 0 2 , p. 87). “ Small, Large tubercles are perforate and crenulate. circular, subcircular, oval or rhomboidal holes or pits of biogenic origin in hard substrates, commonly perpendi R e m a r k s cular to subperpendicular to substrate surface. Excavation The structures described above did not become readily may pass directly through substrate as a penetration, most apparent until the pits were first painted with black food commonly where the substrate is a thin shell, or may end colouring and then whitened with ammonium chloride. within the substrate as a shallow to moderately deep Although these structures are absent in parts of at least depression or short, subcylindrica! pit, commonly with a some specimens, this may be a preservational artifact. depth:width ratio of < 1, with or without a central boss.” The functional significance of these structures, both to the host echinoid and the boring organism, is uncertain, but R e m a r k s they are too close in morphology to the spine-bearing Following the recent paper by T o d d& P a l m e r (2 0 0 2 ), tubercles of the holasteroid test for this to be mere co further examination of the type series of Oichnus exca incidence. This interpretation is strongly reinforced by the vatus D o n o v a n & J a g t , 2 0 0 2 b , by the authors has re example in which a larger primary(?) tubercle is sur vealed new and significant morphological features of the rounded by a circlet of smaller secondary(?) tubercles in substrate that clarify the mode of formation. an arrangement typical ofHemipneustes striatoradiatus. If the interpretation of these structures is correct, it would Oichnus excavatus Donovan & Jagt, 2 0 0 2 b suggest that Oichnus excavatus is not a boring as originally (PI. 1, Figs. 4, 6, 7) interpreted(Donovan & Jagt, 2002b), but more probably an embedment structure that left the ectoderm of the 1993 [unnamed boring inHemipneustes striatoradiatus echinoderm intact. Although debatable, it is most probable (L eske , 1778)]; D efour et al., fig. 3c. that these tubercles did bear spines. This determination 2000 [unnamed boringin Hemipneustes striatoradiatus may seem improbable, as these spines would be numerous (L eske , 1778)]; Jagt , pi. 24, figs. 4-5. and would presumably penetrate the soft tissues of the v* 2002b Oichnus excavatus Donovan & Jagt , pp. 69-73, figs. 2a, c, 3a, b, 4c, 5. trace-forming organism. Counter-intuitively, this may 2002 Oichnus excavatus Donovan & Jagt - D o n o v a n & have been advantageous to the producing organism in P ickerill, p. 86. giving it further anchorage to the test, additional to that 72003 Oichnus excavatus Donovan & Jagt - B l i s s e t t & provided by the concave walls and large central boss Pickerill, pp. 221-223, fig. 2. (discussed by Donovan & Jagt,2002b). Indeed, the func tion of the large central boss is easier to explain if it M a t e r ia l supported spines that were embedded in the soft tissues All pits preserved in tests or test fragments of the holas- of the trace producer. Whatever their origin, these tuber teroid Hemipneustes striatoradiatus (L e s k e , 1778). Spe cles add a unique element to the function of these distinc 122 Stephen DONOVAN & John JAGT
tive pits, although the tubercles themselves must be con D e s c r ip t io n sidered a morphological feature of the host echinoid rather Slender, elongate, straight to arcuate, heterotomously than the trace fossil per se. branched grooved borings of semicircular to more com P ic k e r il l & D o n o v a n (1998) synonymised Tremich- pletely rounded section, preserved in the central third of nus B r e t t , 1985, with Oichnus B r o m l e y based on their the surface of the oyster Agerostrea ungulata. Infestation identical morphology. Subsequently, this ichnotaxo- moderately densely packed. Branches diverge between nomic decision has been supported by some authors and about 45-90°. Borings appear discontinuous in some criticised by others (N ie l s e n & N ie l s e n , 2001, 2002; areas. Associated with circular pits of small diameter D o n o v a n & P ic k e r il l , 2002; T o d d & P a l m e r , 2002; which are particularly common on the sloping, ribbed N ie l s e n et al., in press). sides of the shell (poorly seen in PI. 1, Fig. 5). The diagnosis of Oichnus excavatus was recently emended byB l is s e t t & P ic k e r il l (2003) to include more R e m a r k s conical specimens in a MioceneConus sp. from Jamaica. The main interest of this specimen is in its preservation, The ichnospecific diagnosis given herein defines the type which differs from other reported examples of this ich species and does not embrace the range of morphologies notaxon from these deposits. Further, it is a fine example shown by the Jamaican specimens. The latter may be of preservation of the borings in the shell.Talpina in the deserving of description as a new ichnospecies. type area of the Maastrichtian is otherwise preserved as a cast within a natural mould of the host shell(V o ig t , Ichnogenus Talpina v o n H a g e n o w , 1840 1978). Subsequent to the diagnosis by H ä n t z s c h e l (1975, T y p e ichnospecies p. W133), a variety of shelly substrates have been re Talpina ramosa v o n H a g e n o w , 1840, p. 671, by the corded as infested with fossilTalpina ispp. and Recent subsequent designation ofH ä n t z s c h e l (1962, p. W231). phoronid borings, such as scleractinians, scaphopods, bivalves and crinoids(V o ig t , 1975, 1978). This is not O t h e r ichnospecies unexpected, for the systematic position of the substrate Talpina annulata V o i g t , 1975; T. eduliformis Q u e n - should not be considered a valid ichnotaxobasis for a s t e d t , 1858; T. hirsuta V o i g t , 1975; T. gruberi M a y e r , boring (P ic k e r il l , 1994; B r o m l e y , 1996; P ic k e r il l & 1952; T. scalariformis G h a r e , 1982. D o n o v a n , 1998; D o n o v a n & P ic k e r il l , 2002).
D ia g n o s is Ichnogenus Trypanites M ä g d e f r a u , 1937 (Slightly modified after H ä n t z s c h e l , 1975, p. W 1 3 3 .) (emend.B r o m l e y , 1972) “ Straight to curved tunnel systems in shelly substrates, commonly branched, diameterca. 0.2 mm; numerous T y p e ichnospecies oval or circular openings towards exterior.” Trypanites weisei M ä g d e f r a u , 1937, by monotypy.
R e m a r k s O t h e r ichnospecies The diagnosis ofH ä n t z s c h e l (1975, p. W133) considered Trypanites fimbriatus (S t e p h e n s o n , 1952); T. fosteryeo- that this ichnotaxon was only found ” ... in the rostra of mani C o l e & P a l m e r , 1999; T. solitarius ( v o n H a g e belemnoids ...” As demonstrated herein, other robust, n o w , 1840). shelly substrates were also utilised by these borings (F ü r s ic h et al., 1994). As noted by H ä n t z s c h e l (1975), D ia g n o s is Talpina was originally interpreted as a bryozoan boring, (Slightly modified after B r o m l e y & D ’A l e s s a n d r o , but it is now considered to be a product of the activities of 1987, p. 403.) Single-entrance, cylindrical or subcylind phoronids (V o ig t , 1973, 1975, 1978), the lophophorate rica^ unbranched borings in lithic substrates, having cir horseshoe worms (E m ig & D e M it t e l w ih r , 1999). cular to subcircular cross-section throughout length. The axes of borings may be straight, curved or irregular. Talpina cf. ramosa von Hagenow, 1840 (PI. 1, Fig. 5) R e m a r k s B r o m l e y & D ’A l e s s a n d r o (1987, p. 403) considered M a t e r ia l Trypanites to have a circular cross-section. Specimens A single specimen, NHMM MK 160, preserving a com such as that considered below, with an oval cross-section, plex pattern of borings in the surface of a valve of the fit the revised diagnosis above, yet are morphologically oyster Agerostrea ungulata ( v o n S c h l o t h e im , 1813). distinct from other pouch-shaped borings (compare with Trace fossil not infdled. discussions inB r o m l e y , 1972; B r o m l e y & D ’A l e s s a n d r o , 1987). B r o m l e y & D ’A l e s s a n d r o (1987, p. 404) L o c a l it y a n d h o r iz o n considered straightness versus limited sinuosity ofTry From Ankerpoort-’t Rooth quarry, Meerssen(?) Member, panites ispp. to be unsuitable ichnotaxobases, because Maastricht Formation. Upper Cretaceous; uppermost “ Deviations of a boring from a straight course can be Maastrichtian (Fig. 1). caused by many factors ...” Similarly, too rigid a défini- Ichnology of the type Maastrichtian 123 tion of boring cross-section would eliminate ichnotaxa to aperture boring skewed slightly sideways, subperpen that vary from the diagnosis due to factors controlled in dicular to oyster valve internal surface, showing three part by the nature of the substrate, such as thickness (e.g., infoldings of wall that give it a rope-like appearance. thin versus thick shell) and substrate inhomogeneity (as Distal to the geniculation the shaft of boring (about below). 8 mm in length) shows no such infoldings, and is broad and flattened parallel to the inner surface of the oyster. Trypanites cf. solitarius MÄGDEFRAU, 1937 Main shaft has a constricted and round termination ad (Fig. 2) jacent to the muscle scar of the oyster, which it did not penetrate. This main shaft supports a lozenge-like struc M a t e r ia l ture close to the end, separated from the shaft by a further A single specimen, NHMM LN 7384, within an internal infolding and situated on the side away from the inner mould of a valve of the oyster Rastellum macropterum sensu surface of the valve. Total length c. 11 mm. S t e n z e l , 1971, in which the boring is preserved as a cast. R e m a r k s L o c a l it y a n d h o r iz o n The Mesozoic and Cainozoic fossil record of oysters Upper third of the Nekum Member, Maastricht Forma includes numerous examples in which their commonly tion, CBR-Romontbos quarry, Eben Emael, Bassenge, large, thick shells have been encrusted and penetrated by Liège (Belgium) (for lithostratigraphic section, see J a g t , a diversity of organisms or have in turn overgrown a 1995). Upper Cretaceous; upper Maastrichtian (Fig. 1). variety of organic substrates (T o d d , 1993). Thus, the boring in NHMM LN 7384 is not an unusual phenomenon D e s c r ip t io n per se, but, rather, it is of interest because of its unusual Boring preserved in internal mould of oyster valve as a morphology, which is particularly apparent due to the cast in flint. Boring elliptical in section, with smooth, mode of preservation. unsculptured surface. Aperture conical, round, but asym The boring entered the valve on its internal surface, metrical, opening on inner surface of oyster valve. Boring indicating that the host was a dead oyster shell and most showing strong geniculation close to the aperture. Close probably already disarticulated. The infoldings of the in
Fig. 2 — Cast of Trypanites cf. solitarius (von H agenow , 1840), NHMM LN 7384, preserved in association with an internal mould of a valve o f the oyster Rastellum macropterum sensu S te n z e l. (a) Oyster in plan view, boring right of centre, with base adjacent to muscle scar o f mollusc, (b) Ventral view o f oyster, boring to right and shaped like a recumbent ‘L.’ (c) Dorsal view of oyster, boring to left. All scale bars represent 10 mm. 124 Stephen DONOVAN & John JAGT itial, short shaft perpendicular to the inner valve surface are 407) recognised three forms, A, B and C, ofT. solitarius, interpreted as having been produced in reaction to zones of which the specimen described herein is closest in within the shell structure that were particularly difficult to morphology to their form C. However, because of its penetrate. The broad, conical aperture indicates that the unusual cross-section, NHMM LN 7384 is only included shell calcite was perforated with relative ease and, using tentatively within this ichnospecies. this observation as our guide, it appears that the producing organism penetrated in turn five calcareous layers, sepa rated by four thin, non-calcareous layers. The latter were Acknowledgements undoubtedly the thin organic layers of the shell that, in life, separated the thicker prismatic calcareous layers. That We thank Ron Pickerill (Fredericton, Canada), Paul Taylor (London) these layers were difficult for the producer to penetrate is and Tim Palmer (Aberystwyth, Wales) for comments on aspects of the indicated both by the constrictions of the shaft and also by ichnotaxonomy of the structures discussed herein, Liz Harper (Cam bridge) for shedding light on the organic layers of the bivalve shell and the main shaft distal to the geniculation apparently being Willem Renema (Leiden) for unravelling the complexities of the confined to one shell layer and, in consequence, being electronic Zoological Record. Specimens in Plate 1 were photographed elliptical in section (compare with H a r p e r , 1994). Such using equipment purchased with a grant from the Systematics Associa a sculpture imposed by the substrate represents an unusual tion; photographs in Figure 2 were taken by the Photographic Unit, BMNH. We thank our three distinguished reviewers, Richard Bromley form of xenoglyph ( B r o m le y et al., 1984) (= Fremd (Copenhagen), Mark Wilson (Wooster, Ohio) and Ron Pickerill, for skulptur of V o i g t , 1971). their positive and constructive comments. This is a contribution to Of the ichnospecies of Trypanites, the type T. fimbria S.K.D.’s Nationaal Natuurhistorisch Museum Leiden projects 'Trace fossil studies’ and ‘Palaeontology of the Upper Cretaceous of north tus, andT. fosteryeomani occur perpendicular to the host west Europe,’ and to BMNH project #301 'Palaeoecology of hard substrate. Bromley & D’Alessandro (1987, pp. 406- substrates.’
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Palaeoecological and ichnological significance o f microborings Typescript submitted February 12, 2003 in Quaternary Foraminifera.Palaeontologica Electrónica. Revised typescript received July 15, 2003 126 Stephen DONOVAN & John JAGT
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Figs. 1, 2 Arachnostega gastrochaenae B ertling , 1992, NHMM JJ 12354. 1: Detail of part o f internal mould of right(?) valve of lithophagid bivalve. 2: Complete internal mould o f same valve. Fig. 3 - Arachnostega gastrochaenae B e r tlin g , 1992, NHMM MK 367, detail of part of internal mould o f patellid gastropod showing sparsely developed burrows. Figs. 4, 6, - Oichnus excavatus D onovan & Jagt , 2002b, NHMM MK 4689, circular pits in a test of the holasteroid echinoid Hemipneustes striatoradiatus (L esk e). Paratypes unless stated otherwise. 4: Pit showing tubercles on wall and side of central boss. 6: Three pits (holotype arrowed); confluent pair show tubercles on central boss and floor of holotype. 7; Two strongly tuberculated pits. Right example has two sizes of tubercles on central boss and bevelled lip o f pit. Fig. 5 - Talpina cf. ramosa von Hagenow , 1840, NHMM MK 160, borings in shell of oysterAgerostrea ungulata (von Schlotheim ), Nekami, Bemelen, southern Limburg, Meerssen(?) Member, Maastricht Formation.
All specimens painted with black food colouring and subsequently coated with ammonium chloride sublimate. All scale bars represent 10 mm. Ichnology of the type Maastrichtian 127
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