The Maastrichtian of the Type Area (Upper Cretaceous, the Netherlands and Belgium) by Stephen K
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
Donovan, S.K. & Jagt, 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 l'Institut royal 1840 généralement conservé sous forme de moulages naturels dans des des Sciences naturelles de Belgique, Sciences de la Terre, 74: 119-127, coquilles décalcifiées, se présente sous forme de cavités dans l'huître 2 figs., 1 pl., Bruxelles-Brussel, March 31, 2004. - ISSN 0374-6291. Agerostrea ungulata (von Schlotheim, 1813). Trvpanites cf. solitarius Màgdefrau, 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 Mots-clefs: Cretaceous) are described, and their morphology and palaeoecological Terriers, forages, "enfouissement", Maastrichtien, Pays- significance discussed; a fourth ichnotaxon shows an unusual mode of Bas, Belgique. préservation. The infill of an articulated lithophagid bivalve (Meerssen Member, Maastricht Formation) preserves the distinctive firmground burrow Arachnostega gastrochaenae Bertling, 1992. This is the first Introduction report of this trace fossil, probably generated by polychaetes, from the Cretaceous of northem 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 element of the biota of the Maastrichtian are recognised for the first time within the pits of Oichnus excavatus (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) (Dortangs, 1998; The producing organism may have obtained a firmer attachment to the Donovan & test by having abundant spines of the host echinoid embedded in its soft Jagt, 2002a; Jagt, 2003) (Fig. 1). The present tissues. The boring Talpina cf. ramosa von flagenow, 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). Trvpanites cf. the various members included in the Maastricht Formation. solitarius Màgdefrau, 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 Rasteüum macropterum logically distinct ichnofossils fforn this unit, each of which sensu Stenzel, 1971 (Nekum Member, Maastricht Formation). shows unusual features related to préservation, 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 FIÀntzschel (1975). The philosophy of open nomencla¬ Résumé ture follows Bengtson (1988). Annotations within syno- nymy lists follow Matthews (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 Netherlands tion paléoécologique sont discutées. Un quatrième ichnotaxon montre un (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 Ichnogenus Arachnostega Bertling, 1992 montre un autre exemple. Des tubercules ont été observés pour la première fois dans les cavités d'Oichnus excavatus Donovan & Jagt, Type ichnospecies 2002b (Membre de Meerssen, Formation de Maastricht) indiquant qu'elles résultent probablement d'un "enfouissement" plutôt que d'un Arachnostega gastrochaenae Bertling, 1992, p. 179, by creusement. L'organisme responsable peut avoir été plus fermement fixé original désignation. The only nominal ichnospecies. 120 Stephen DONOVAN & John JAGT
1994 Arachnostega gastrochaenae Bertling - fürsich et al., pp. 146, 161, pl. 3, figs. 1, 2, 4. ? 2001 ftrac/yzcwtega-Wilson & Taylor, p. 26, pl. 2, fig. 1.
Material Two specimens. An incomplete internai mould of an indeterminate lithophagid bivalve, NF1MM JJ 12354, with A. gastrochaenae apparent within the moulds of both valves (Pl. 1, Figs. 1, 2). The internai mould of the bivalve, preserved in a medium- to coarse-grained bio- calcarenite, is broken both anteriorly and posteriorly, preserving little detail of the internai surface of the mol- lusc shell except for some coarse growth lines. The second specimen, NHMM MK 367, is an internai mould of an indeterminate patel lid limpet of the type described by Kaunhowen (1898, p. 15, pl. 1, figs. 3-4), with A. gastrochaenae weakly developed (Pl. 1, Fig. 3). Fig. 1 — Outline map of study area (redrawn and simplified after Jagt, 1999, fig. 1), showing political boundaries Locality and horizon (dashed lines), rivers and canals (solid lines) and the NFIMM JJ 12354 is from ENCI-Maastricht BV quarry, city of Maastricht (M). = Key to localities: 1 ENC1- 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 trichtian (NI), Belgium (B) and Germany (D) shows the ap- (for simple lithostratigraphic section, see Jagt et proximate position of the main map (box). al., 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). Diagnosis Diagnosis (From Bertling, 1992, p. 179.) "Irregular elongate and net-like burrows in sediment ftlls of shells. Visible on the (After Bertling, 1992, p. 180). "Ramified burrows on the surface of internai moulds with an oval surface of internai moulds. The size of the meshwork may cross-section, which increases vary from microns to centimetres, depending on the shell- 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. Latéral burrows mostly have a smaller Remarks diameter. Polygonal areas on the surface of internai moulds Arachnostega is a distinctive network of channels that is may be produced by the unification of ramified burrows." superfïcially reminiscent of structures such as the clionid sponge boring Entobia Bronn, 1838, although lacking Description Based swollen chambers and not penetrating a calcareous mainly on NJFMM JJ 12354. Burrows more inten- shell or limestone substrate. That Arachnostega itself is sely developed adjacent to the right(?) valve (Pl. 1, Figs. 1, not a boring was only determined by the astuteness of 2) than left(?). Burrows best seen on surface of moulds, Bertling (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 (Pl. 1, Fig. 2), revealing less common ground settings within sediment-filled shells of the in- burrows below. Burrows of incomplete circular or ellipti- cal faunal bivalve Mya sp. within the North Sea Basin, section, preserved as grooves with overhanging edges and not generated by more than one taxon of polychaete annelid infilled, individual burrows varying from c. 0.1 to (Reineck, 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 rows(?) may be a taphonomic artifact. Larger diameter Jurassic of northern Europe (Bertling, 1992; Fürsich et burrows forming reticulate pattem at surface of mould. al., 1994) and the Cretaceous of the Middle East (Wilson & Taylor, 2001). Remarks This is the first reported occurrence of this distinctive, but Arachnostega gastrochaenae Bertling, 1992 cryptic, ichnotaxon from the Maastrichtian of the type (Pl. 1, Figs. 1-3) area. Wilson & Taylor (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- Todd, p. 417. sual mode of occurrence of A. gastrochaenae within in- Ichnology of the type Maastrichtian 121 ftlled shells and borings prevents its identification except cimens considered herein include the holotype pit in within mouldic material, so it is probably more widespread NHMM MK 4689 (Pl. 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 in Fürsich et al., 1994). (Pl. 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 intemal mould, where movement would have been constrained by Locality and horizon 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. Jagt, 2002b, fig. 1 ). Upper Cretaceous; uppermost Maas¬ trichtian (Fig. 1). Ichnogenus Oichnus Bromley, 1981 Diagnosis
Type ichnospecies (Slightly revised after Donovan & Jagt, 2002b, p. 69.) Oichnus simplex Bromley, 1981, p. 60, by original dés¬ Circular to elliptical, non-penetrative Oichnus, almost ignation. invariably with a broad, high, raised central boss. Aper- ture of pit overhanging and walls concave. Other ichnospecies Description Oichnus asperus Nielsen & Nielsen, 2001 ; O. coronatus Nielsen & Nielsen, 2001; O. excavatus Donovan & (Further to features described in Donovan & Jagt, Jagt, 2002b; O. gradatus Nielsen & Nielsen, 2001; 2002b.) All surfaces of pits bearing sculpture of irregu- O. ovalis Bromley, 1993; O. paraboloides Bromley, larly distributed to well-ordered tubercles, commonly 1981. very small (granular), although may be present in two sizes (Pl. 1, figs 4, 6, 7). In at least one shallow pit, small Diagnosis tubercles are arrayed around larger tubercles in a circle. (After Donovan & Pickerill, 2002, 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- Remarks The structures described above did not become readily cular to subperpendicular to substrate surface. Excavation may pass directly through substrate as a pénétration, 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 some dépression or short, subcylindrical pit, commonly with a specimens, this may be a preservational artifact. The functional depth:width ratio of ^ 1, with or without a central boss." significance of these structures, both to the host echinoid and the boring organism, is uncertain, but Remarks they are too close in morphology to the spine-bearing tubercles of the holasteroid test for this to be mere co¬ Following the recent paper by Todd & Palmer (2002), further examination of the type series of Oichnus exca¬ ïncidence. This interprétation is strongly reinforced by the vatus Donovan & Jagt, 2002b, by the authors has re- example in which a larger primary(?) tubercle is sur- rounded vealed new and significant morphological features of the by a circlet of smaller secondary(?) tubercles in substrate that clarify the mode of formation. an arrangement typical ofHemipneustes striatoradiatus. If the inteipretation of these structures is correct, it would Oichnus excavatus Donovan & Jagt, 2002b suggest that Oichnus excavatus is not a boring as originally (Pl. 1, Figs. 4, 6, 7) interpreted (Donovan & Jagt, 2002b), but more probably an embedment structure that left the ectoderm of the 1993 [unnamed boring in Hemipneustes striatoradiatus echinoderm intact. Although debatable, it is most probable (Leske, 1778)]; Defour et al., fig. 3c. that these tubercles did bear spines. This détermination 2000 [unnamed boring in Hemipneustes striatoradiatus may seem improbable, as these spines would be numerous (Leske, 1778)]; Jagt, pl. 24, figs. 4-5. and would presumably penetrate the soft tissues of the v* 2002b Oichnus excavatus Donovan & Jagt, pp. 69-73, trace-forming organism. Counter-intuitively, this may figs. 2a, c, 3a, b, 4c, 5. have been advantageous to the producing organism in 2002 Oichnus excavatus Donovan & Jagt - Donovan & giving it further anchorage to the test, additional to Pickerill, p. 86. that 72003 Oichnus excavatus Donovan & Jagt - Blissett & provided by the concave walls and large central boss Pickerill, pp. 221-223, fig. 2. (discussed by Donovan & Jagt, 2002b). Indeed, the ftinc- tion of the large central boss is easier to explain if it Material supported spines that were embedded in the soft tissues of the trace producer. Whatever All pits preserved in tests or test fragments of the holas- their origin, these tuber¬ cles add a teroid Hemipneustes striatoradiatus (Leske, 1778). Spe¬ unique element to the function of these distinc- 122 Stephen DONOVAN & John JAGT
tive pits, although the tubercles themselves must be con- Description 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- Pickerill & Donovan (1998) synonymised Tremich- pletely rounded section, preserved in the central third of nus Brett, 1985, with Oichnus Bromley based on their the surface of the oyster Agerostrea ungulata. Infestation identical morphology. Subsequently, this ichnotaxo- moderately densely packed. Branches diverge between nomic décision has been supported by some authors and about 45-90°. Borings appear discontinuous in some criticised by others (Nielsen & Nielsen, 2001, 2002; areas. Associated with circular pits of small diameter Donovan & Pickerill, are 2002; Todd & Palmer, 2002; which particularly common on the sloping, ribbed Nielsen et al., in press). sides of the shell (poorly seen in Pl. 1, Fig. 5). The diagnosis of Oichnus excavatus was recently emended by Blissett & Pickerill (2003) to include more Remarks conical specimens in a Miocene Conus sp. from Jamaica. The main interest of this specimen is in its préservation, 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 of specimens. The latter may be préservation 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 (Voigt, Ichnogenus Talpina von Hagenow, 1840 1978). Subséquent to the diagnosis by Hàntzschel (1975, Type ichnospecies p. W133), a variety of shelly substrates have been re- Talpina ramosa von Hagenow, 1840, as p. 671, by the corded infested with fossil Talpina ispp. and Recent subséquent désignation of Hàntzschel (1962, p. W231). phoronid borings, such as scleractinians, scaphopods, bivalves and crinoids (Voigt, 1975, 1978). This is not Other ichnospecies unexpected, for the systematic position of the substrate Talpina annulata Voigt, 1975; T. not eduliformis Quen- should be considered a valid ichnotaxobasis for a stedt, 1858; T. hirsuta Voigt, 1975; T. gruberi Mayer, boring (Pickerill, 1994; Bromley, 1996; Pickerill & 1952; T. scalariformis Ghare, 1982. Donovan, 1998; Donovan & Pickerill, 2002). Diagnosis Ichnogenus Trypanites Màgdefrau, 1937 (Slightly modified after Hàntzschel, 1975, p. W133.) (emend. Bromley, 1972) "Straight to curved tunnel Systems in shelly substrates, commonly branched, diameter ca. 0.2 mm; numerous Type ichnospecies oval or circular openings towards exterior." Trypanites weisei Màgdefrau, 1937, by monotypy. Remarks Other ichnospecies The diagnosis of Hàntzschel (1975, p. W133) considered Trypanites fimbriatus (Stephenson, that this 1952); T. fosteryeo- ichnotaxon was only found "... in the rostra of mani Cole & Palmer, 1999; T. solitarius (von Hage¬ belemnoids ..." As demonstrated herein, other robust, now, 1840). shelly substrates were also utilised by these borings (Fürsich et al., 1994). As noted by Hàntzschel (1975), Diagnosis Talpina was original ly interpreted as a bryozoan boring, (Slightly modified after Bromley & but it is D'Alessandro, now considered to be a product of the activities of 1987, p. 403.) Single-entrance, cylindrical or subcylind- phoronids (Voigt, 1973, 1975, 1978), the lophophorate rical, unbranched borings in lithic substrates, horseshoe worms having cir¬ (Emig & De Mittelwihr, 1999). cular to subcircular cross-section throughout length. The axes of borings may be straight, curved or irregular. Talpina cf. ramosa von Hagenow, 1840 (Pl. 1, Fig. 5) Remarks Bromley & D'Alessandro Material (1987, p. 403) considered Trypanites to have a circular cross-section. A Specimens single specimen, NHMM MK 160, such as that preserving a com¬ 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 (von Schlotheim, 1813). distinct from other pouch-shaped borings Trace fossil not infilled. (compare with discussions in Bromley, 1972; Bromley & D'Alessan¬ dro, 1987). Bromley & D'Alessandro Locality and horizon (1987, p. 404) considered straightness versus limited From Ankerpoort-'t Rooth sinuosity of Try¬ quarry, Meerssen(?) Member, panites ispp. to be unsuitable Maastricht Formation. ichnotaxobases, because Upper Cretaceous; uppermost "Déviations of a Maastrichtian boring from a straight course can be (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 internai 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 Màgdefrau, 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¬ Material ture close to the end, separated from the shaft by a further A single specimen, NHMM LN 7384, within an internai infolding and situated on the side away from the inner mould of a valve ofthe oyster Rastellum macropterum sensu surface of the valve. Total length c. 11 mm. Stenzel, 1971, in which the boring is preserved as a cast. Remarks
Locality and horizon 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 Jagt, a diversity of organisms or have in turn overgrown a 1995). Upper Cretaceous; upper Maastrichtian (Fig. 1). variety of organic substrates (Todd, 1993). Thus, the boring in NFIMM LN 7384 is not an unusual phenomenon Description per se, but, rather, it is of interest because of its unusual Boring preserved in internai 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 préservation. unsculptured surface. Aperture conical, round, but asym¬ The boring entered the valve on its internai 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 Hagenow, 1840), NHMM LN 7384, preserved in association with an internai mould of a valve of the oyster Rastellum macropterum sensu Stenzel. (a) Oyster in plan view, boring right of centre, with base adjacent to muscle scar of mollusc. (b) Ventral view of 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, of T. 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¬ the main shaft distal to the geniculation apparently being bridge) for shedding light on the organic layers of the bivalve shell and Willem Renema (Leiden) for unravelling the complexities of the confined to one shell layer and, in conséquence, being electronic Zoological Record. Specimens in Plate 1 were photographed elliptical in section (compare with Harper, 1994). Such using equipment purchased with a grant from the Systematics Associa¬ a tion; in were sculpture imposed by the substrate represents an unusual photographs Figure 2 taken by the Photographie Unit, BMNH. We thank our three fonn of distinguished reviewers, Richard Bromley xenoglyph (Bromley et al., 1984) (= Fremd- (Copenhagen), Mark Wilson (Wooster, Ohio) and Ron Pickerill, for skulptur of Voigt, 1971). their positive and constructive comments. This is a contribution to S.K.D.'s Of the ichnospecies of Trypanites, the type T. fimbria- Nationaal Natuurhistorisch Museum Leiden projects 'Trace fossil studies' and 'Palaeontology of the Upper Cretaceous of north- tus, and T. fosteryeomani occur perpendicular to the host west Europe,' and to BMNH project #301 'Palaeoecology of hard substrate. Bromley & D'Alessandro (1987, pp. 406- substrates. '
References
Bengtson, P., 1988. Open nomenclature. Palaeontology, 31: Cole, A.R. & Palmer, T.J., 1999. Middle Jurassic worm 223-227. borings, and a new giant ichnospecies of Trypanites from the Bertling, M., 1992. Arachnostega n. ichnog. - burrowing Bajocian/Dinantian unconformity, southem England. Pro- traces in internai moulds of boring bivalves (Late Jurassic, ceedings of the Geologists' Association, 110: 203-209. Northern Germany). Palàontologische Zeitschrift, 66: 177-185. Defour, E., Geussens, T., Indeherberge, L. & Strijbos, V., 1993. van Blissett, D.J. & Pickerill, R.K., 2003. Oichnus excavatus Vormvariaties Hemipneustes striatoradiatus en Donovan & Jagt, 2002, from the Moneague Formation, White Hemiaster prunella uit het Boven-Krijt van Limburg. LIKONA Limestone Group, Jamaica. Caribbean Journal ofScience, 39: (Limburgse Koepel voor Natuurstudie), Jaarboek 1993: 7-14. 221-223. Donovan, S.K. & Jagt, J.W.M., 2002a. Ichnology of the type Brett, C.E., 1985. Tremichnus: a new ichnogenus of circular- area of the Maastrichtian Stage (Upper Cretaceous): burrowing parabolic pits in fossil echinoderms. Journal of Paleontology, and boring immediately prior to the K/T boundary event. 59: 625-635. Palaeontological Association Newsletter, #51: 91. Bromley, R.G., 1972. On some ichnotaxa in hard substrates, Donovan, S.K. & Jagt, J.W.M., 2002b. Oichnus Bromley with a redéfinition of Trypanites Mâgdefrau. Palàontologische borings in the irregular echinoid Hemipneustes Agassiz from Zeitschrift, 46: 93-98. the type Maastrichtian (Upper Cretaceous, The Netherlands and Bromley, R.G., 1981. Concepts in ichnotaxonomy illustrated Belgium). lchnos, 9: 67-74. by small round holes in shells. Acta Geológica Hispànica, 16: Donovan, S.K. & Pickerill, R.K., 2002. Pattern versus pro- 55-64. cess or informative versus uninformative ichnotaxonomy: reply to Todd and Palmer. Bromley, R.G., 1993. Prédation habits of octopus past and Ichnos, 9: 85-87. present and a new ichnospecies, Oichnus ovalis. Bulletin of Dortangs, R.W., 1998. Sporenfossielen. In: Jagt, J.W.M., the Geological Society ofDenmark, 40: 167-173. Leloux, J. & Dhondt, A.V. (Editors), Fossielen van de St. Bromley, R.G. 1996. Trace Fossils: Biology, Taphonomy and Pietersberg. Grondboor & Hamer, 52 (4/5): 150-151. Applications (2nd édition). Chapman and Hall, London, Emig, C.C. & Mittelwihr, C. de, 1999. What is a phoronid? xvi + 361 pp.
Plate 1
Figs. 1,2 — Arachnostega gastrochaenae Bertling, 1992, NHMM JJ 12354. 1: Detail of part of internai mould of right(?) valve of lithophagid bivalve. 2: Complete internai mould of same valve. Fig. 3 — Arachnostega gastrochaenae Bertling, 1992, NHMM MK 367, detail of part of internai mould of patellid gastropod showing sparsely developed burrows. Figs. 4, 6, 7 — Oichnus excavatus Donovan & Jagt, 2002b, NHMM MK 4689, circular pits in a test of the holasteroid echinoid Hemipneustes striatoradiatus (Leske). 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 of pit. Fig. 5 — Talpina cf. ramosa von Hagenow, 1840, NHMM MK 160, borings in shell of oyster Agerostrea 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 125
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Nationaal Natuurhistorisch Museum Mayer, G., 1952. Lebensspuren von Bohrorganismen aus dem Postbus 9517, NL-2300 RA Leiden unteren Hauptmuschelkalk des Kraichgaues. Neues Jahrbuch The Netherlands für Geologie und Palaontologie, Monatshefte, 1952: 450-456. E-mail: [email protected] Nielsen, J.K. & Nielsen, K.S.S., 2002. Pattern versus process or informative versus ininformative (sic) ichnotaxonomy: reply John W. M. Jagt to Todd and Palmer. Ichnos, 9: 83-84. Natuurhistorisch Museum Maastricht Nielsen, K.S.S. & Nielsen, J.K., 2001. Bioerosion in Pliocene de Bosquetplein 6 to late Holocene tests of benthic and planktonic foraminiferans, NL-6211 KJ Maastricht with a revision of the ichnogenera Oichnus and Tremichnus. The Netherlands Ichnos, 8: 99-116. E-mail: [email protected] Nielsen, K.S.S., Nielsen, J.K. & Bromley, R.G. (in press). Palaeoecological and ichnological significance of microborings Typescript submitted February 12, 2003 in Quaternary Foraminifera. Palaeontologica Electronica. Revised typescript received July 15, 2003 Plate 1