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Stratigraphical ranges of tegulated inoceramid bivalves in the type area of the Maastrichtian Stage (Belgium, the Netherlands)
Accepted Manuscript Stratigraphical ranges of tegulated inoceramid bivalves in the type area of the Maastrichtian Stage (Belgium, the Netherlands) John W.M. Jagt, Elena A. Jagt-Yazykova PII:
S0195-6671(17)30157-X
DOI:
10.1016/j.cretres.2017.05.022
Reference:
YCRES 3619
To appear in:
Cretaceous Research
Received Date: 1 April 2017 Revised Date:
20 May 2017
Accepted Date: 21 May 2017
Please cite this article as: Jagt, J.W.M., Jagt-Yazykova, E.A., Stratigraphical ranges of tegulated inoceramid bivalves in the type area of the Maastrichtian Stage (Belgium, the Netherlands), Cretaceous Research (2017), doi: 10.1016/j.cretres.2017.05.022. This is a PDF file of an unedited manuscript that has been accepted for publication. As a service to our customers we are providing this early version of the manuscript. The manuscript will undergo copyediting, typesetting, and review of the resulting proof before it is published in its final form. Please note that during the production process errors may be discovered which could affect the content, and all legal disclaimers that apply to the journal pertain.
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Stratigraphical ranges of tegulated inoceramid bivalves in the type area of
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the Maastrichtian Stage (Belgium, the Netherlands)
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John W.M. Jagt a,*, Elena A. Jagt-Yazykova b
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a
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b
Uniwersytet Opolski, Katedra Biosystematyki, Pracownia Paleobiologii, ul. Oleska 22, 45-052
Opole, Poland
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Natuurhistorisch Museum Maastricht, De Bosquetplein 6-7, 6211 KJ Maastricht, the Netherlands
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E-mail address: [email protected] (J.W.M. Jagt).
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ABSTRACT
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Recently collected material, in fine- to medium-grained biocalcarenites and in coeval flint
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nodules, of the tegulated inoceramid taxa Spyridoceramus tegulatus (von Hagenow, 1842)
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and Tenuipteria argentea (Conrad, 1858) from various members of the Gulpen Formation and
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the overlying Maastricht Formation allows their stratigraphical ranges to be refined. It is
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concluded that the ranges correspond closely to those in east-central Poland (Vistula [Wisła]
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River valley region), where S. tegulatus occurs in the lower Maastrichtian and lower upper
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Maastrichtian (Endocostea typica, Trochoceramus radiosus and ‘Inoceramus’ ianjonaensis
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inoceramid zones, or Belemnella occidentalis and Belemnitella junior cephalopod zones). In
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contrast, Tenuipteria argentea appears to be confined to the overlying zone of Belemnella
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(Neobelemnella) kazimiroviensis (Skołozdrówna, 1932) and Belemnella (N.) skolozdrownae
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(Kongiel, 1962) and there is no overlap in the range of both species. In the type area of the
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ACCEPTED MANUSCRIPT Maastrichtian Stage, S. tegulatus ranges from the Vijlen Member (interval 3) of the Gulpen
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Formation to the top of the Nekum Member of the Maastricht Formation, while T. argentea is
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restricted to the upper Meerssen Member (subunits IVf-6 and IVf-7, with an acme in the
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former), straddling the Cretaceous-Paleogene (K/Pg) boundary.
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Keywords:
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Mollusca
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Inoceramidae
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Cretaceous
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Stratigraphy
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Correlations
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Northwest Europe
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1. Introduction
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In the extended type area of the Maastrichtian Stage (Fig. 1), inoceramid bivalves generally are rather uncommon and often of mediocre or poor preservation. Notable
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exceptions include assemblages from the Vaals Formation (lower lower Campanian; see Van
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der Weijden, 1943; Sornay, 1982; Walaszczyk and Dhondt, 2005), the Zeven Wegen Member
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(Gulpen Formation; lower upper Campanian) and intervals 3–6 of the Vijlen Member (Gulpen
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Formation, mid- to lower upper Maastrichtian; see Walaszczyk et al., 2010). The youngest
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true inoceramids, ‘Inoceramus’ ianjonaensis morgani Sornay, 1973, Cataceramus
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ghadamesensis (Tröger, in Tröger and Röhlich, 1981) and Endocostea coxi (Reyment, 1955),
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are known from interval 6 of the Vijlen Member, associated with a single tegulated taxon,
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Spyridoceramus tegulatus. Higher upsection, from the Lanaye Member (Gulpen Formation)
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onwards, only thin-shelled, and mostly ill-preserved, tegulated forms remain, occasionally
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occurring in coquina-like concentrations.
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The first tegulated inoceramid to have been recorded from the Maastrichtian type area, although not recognised as such, was Avicula geulemensis Vogel, 1895 (Fig. 2). Of this
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species, now referred to as Tenuipteria argentea, an extensive material has been collected at a
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number of localities in the Dutch and Belgian provinces of Limburg over recent years. The
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same holds true for Spyridoceramus tegulatus, so that the stratigraphical ranges of these taxa
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can now be determined in more detail.
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2. Previous studies
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As documented by Dhondt (1983, 1992), tegulated inoceramids of the genera Spyridoceramus Cox, in Moore, 1969 and Tenuipteria Stephenson, 1955 appear restricted to
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North America (Western Interior, Gulf and Atlantic coasts) and western Eurasia. Dhondt
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(1983) noted that the equivalve S. tegulatus ranged from the lower Maastrichtian to the lower
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upper Maastrichtian, with records from the Western Interior, the European white chalk facies,
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the Russian Platform and central Asia. The inequivalve Tenuipteria argentea was shown to be
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restricted to the upper Maastrichtian, with records from the North American Gulf and Atlantic
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coasts into the European white chalk facies, the Russian Platform and central Asia. Amongst
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named forms, equivalve taxa (see Dhondt, 1983, p. 45) include ‘Inoceramus’ tegulatus,
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Avicula? fibrosa Meek and Hayden, 1856 and ‘I.’ caucasicus Dobrov, 1951, while
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inequivalve taxa comprise ‘Inoceramus’ argenteus, ‘Inoceramus’ dobrovi Jeletzky, in
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Jeletzky and Clemens, 1965 and Avicula geulemensis. This distinction between inequivalve
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and equivalve forms has been referred to in the literature as ‘Avicula-like’ or ‘submytiloid to
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subquadrate’ (i.e., S. tegulatus) and ‘Pholadomya-like’ (i.e., T. argentea). In general,
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ornament in these two taxa is quite similar, although there is a certain range of variation.
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Dhondt (1982, 1983) noted an overlap in the stratigraphical ranges of both taxa in the lower
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part of the Belemnitella junior cephalopod Zone in northern Germany (see also Tröger et al.,
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2001, fig. 13), albeit on the basis of rather poorly preserved material. Abdel-Gawad (1986) noted that there had been some confusion between the two taxa
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amongst previous workers, mostly on account of close similarities in the tile-like (tegulated)
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ornament, taphonomic deformation and the general lack of complete, articulated specimens,
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mainly of S. tegulatus. Unlike Dhondt (1983), Abdel-Gawad did not note any overlap in
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ranges; a notion corroborated in the present paper. In fact, Jagt (1996, 1999) suggested that T.
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argentea was confined to the upper Meerssen Member and that the range of this taxon in the
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Maastrichtian type area corresponded to that in east-central Poland.
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Recently, Walaszczyk (in Walaszczyk et al., 2016, p. 317) has noted that European inoceramid faunas of Campanian and Maastrichtian age were representative of a uniform
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Euramerican Biogeographic Region, and that the final extinction of true inoceramids occurred
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during the early late Maastrichtian (see above). The late Maastrichtian species, referred to as
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tegulated inoceramids (or inoceramoids, sensu Tröger et al., 2001), had invariably been
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regarded to be distinct from genuine inoceramids on features of their ligamental plates.
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However, the loss of regularity in ligament pits had also been seen in a few late Campanian
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and early Maastrichtian inoceramids (Zonova, 1980; Walaszczyk, 2004), so that this specific
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character could not be used for suprageneric classification of such forms (Walaszczyk in
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Walaszczyk et al., 2016).
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These data are here corroborated, and the stratigraphical ranges of S. tegulatus and T.
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argentea in the type area of the Maastrichtian Stage refined. For more details on the
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palaeogeography and lithostratigraphy of the Maastrichtian type area within northwest
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Europe, reference is made to Jagt (1999) and Voigt et al. (2008).
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3. Systematic palaeontology Abbreviations. To denote the repositories of specimens illustrated and/or referred to in the text, the following abbreviations are used: IRScNB, Institut royal des Sciences naturelles
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de Belgique, Brussels, Belgium; NCB, Naturalis Biodiversity Center, Leiden, the
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Netherlands; NHMM, Natuurhistorisch Museum Maastricht, Maastricht, the Netherlands.
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Family Inoceramidae Giebel, 1852
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Genus Spyridoceramus Cox in Moore, 1969 (ex Heinz, 1932, nomen nudum)
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Type species. Inoceramus tegulatus von Hagenow, 1842 (non Ødum, 1922), by original
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designation.
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Spyridoceramus tegulatus (von Hagenow, 1842)
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Figs 3A–K, 4A–K, 5A–D.
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*1842 Inoceramus tegulatus von Hagenow, p. 559.
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*1856 Avicula? fibrosa Meek and Hayden, p. 86.
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*1951 Inoceramus caucasicus Dobrov, p. 167, pl. 2, fig. 2.
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1959 Inoceramus caucasicus Dobrov; Dobrov and Pavlova in Moskvin, p. 150, pl. 19, fig.
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4a–b.
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1970a Tenuipteria fibrosa (Meek & Hayden); Speden, p. 62, pl. 8, figs 1–18; pl. 9, figs 1–16.
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1979 Tenuipteria geulemensis (F. Vogel, 1895); Dhondt, p. 142 (pars), pl. 1, figs 1, 3–4 (non
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figs 2, 5–6 = Tenuipteria argentea). 1982 Tenuipteria geulemensis (Vogel, 1895); Dhondt, p. 77 (pars), pl. 1, figs 1–4, ?5–8 (non figs 9–11 = Tenuipteria argentea).
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1986 Spyridoceramus tegulatus (v. Hagenow, 1842); Abdel-Gawad, p. 146, pl. 31, figs 1–2 (with additional synonymy). 2001 Spyridoceramus cf. tegulatus (Hagenow 1842); Tröger et al., p. 156, text-fig. 12; pl. 1, fig. 6; pl. 3, fig. 2. 2010 Spyridoceramus tegulatus (von Hagenow, 1842); Walaszczyk et al., p. 162, fig. 9/1–8
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(with additional synonymy).
2016 Spyridoceramus tegulatus (v. Hagenow, 1842); Machalski and Malchyk, p. 575, fig. 3D.
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Material. About 50 specimens, preserved both in flint nodules and in fine- to medium-grained
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biocalcarenites, including IRScNB TCMI 10165, IRScNB TCMI 10167, NHMM 2006 083,
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NHMM 2010 127a–b, NHMM 2016 048, NHMM 2017 003, NHMM 2017 005, NHMM
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2017 006a–b, NHMM 2017 007, NHMM JJ 6188a–c, NHMM JJ 6355, NHMM JJ 7011,
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NHMM JJ 11853, NHMM JJ 11965, NHMM JJ 13540, NHMM JJ 14238, NHMM JJ 14508,
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NHMM JJ 14803a–b, NHMM JJ 15029 and NHMM JJ 15895.
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Description. Except for flint-preserved material (e.g., Figs 3A–C, F, H, J, 4B, J, 5A), all
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specimens have suffered from sediment compaction and are fragmentary to varying degrees.
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A number of specimens represent small pieces of shell only, but can be recognised on account
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of their specific ornament. At certain levels, shells occur in clumps, in a coquina-like fashion,
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occasionally as articulated, albeit heavily deformed individuals (e.g., Figs 3K, 4C). For this
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reason, a full description cannot be offered, but specific features of the various shells are
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indicated below.
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Remarks. On the basis of material of S. tegulatus from the Vijlen Member (Gulpen
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Formation) in the Aachen (Germany) and Lixhe-Haccourt (Belgium) areas, Walaszczyk et al.
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(2010, p. 163, figs 2, 9) distinguished three formae, A–C. Their ‘forma A’, occurring in
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interval 3 (Trochoceramus radiosus inoceramid Zone) of that member, is axially elongated
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and slender, with an ornament consisting predominantly of concentric rugae. Radial elements
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ACCEPTED MANUSCRIPT of the ornament comprise discontinuous ribs that form rows of nodes where they and
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concentric rugae intersect. In shell shape and ornament, ‘forma A’ is close to the North
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American Tenuipteria fibrosa, in particular to material from the Fox Hills Formation of South
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Dakota as illustrated by Speden (1970a, b). The lectotype of S. tegulatus from the lower
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Maastrichtian of Rügen (northeast Germany) probably represents the same form (see
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Wolansky, 1933, pl. 5, fig. 6).
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‘Forma B’ is a broader morphotype in which ornament consists nearly exclusively of concentric rugae. This is found in the upper part of interval 5 and in the lower part of interval
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6 (‘Inoceramus’ ianjonaensis inoceramid Zone) of the Vijlen Member, and comes close to
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Inoceramus caucasicus var. ravni Dobrov, 1951. Lastly, confined to the lower part of interval
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6 is ‘forma C’, with very strong radial ornament, in particular anteriorly. This bears a strong
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resemblance to Inoceramus caucasicus var. fistulata Dobrov, 1951 and to one of Wolansky’s
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(1933, pl. 4, fig. 5) syntypes of S. tegulatus.
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be fully assessed in view of the paucity of well-preserved specimens which has so far
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precluded a stratophenetic analysis of the European material. Specimens from the upper
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Lanaye Member (Gulpen Formation) and the Valkenburg, Schiepersberg, Emael and Nekum
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members (Maastricht Formation) generally are of medium size. They all show well-marked
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concentric and radial ornament, covering the anterior and median parts of the shell (where this
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can be determined) or with radial plicae more prominent in the anterior part of the shell.
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Where concentric and radial elements intersect, quadrate to rectangular nodes are seen, best
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developed in the anterior and median parts of the shell. The strong radial ornament (i.e., deep
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grooves anteriorly) of ‘forma C’ is mostly lacking, although a large-sized, flint-preserved left
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valve (Figs 3H, 5A) shows a number of shallow furrows that continue to the ventral margin in
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the anterior portion of the shell. In overall proportions, this individual resembles a specimen
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ACCEPTED MANUSCRIPT of Inoceramus fibrosus from Canada illustrated by Jeletzky (1962, pl. 141, fig. 6), although
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radial plicae are much more prominent, and ornament differs in details. All in all, there is a
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close match between specimens that we have studied and material from the upper
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Maastrichtian of North America (Speden, 1970a, pl. 9, as Tenuipteria fibrosa) and from the
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‘Kunrade Limestone’ (now Kunrade Formation) in the study area as illustrated by Dhondt
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(1979, pl. 1, fig. 3, in particular).
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It would appear that, in shell shape and ornament, the specimens studied by us are uniform (Figs 3A–C, E–K; 4A–C, J–K; 5A, C), although a certain ecophenotypic impact on
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shell morphology may be implied. Where shell (< 1 mm in thickness) material is available,
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the colour is orange brown to dark brown.
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Occurrence. The present species ranges from the lowermost Maastrichtian (upper part of the
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‘Inoceramus’ redbirdensis inoceramid Zone) into the lower upper Maastrichtian (high into the
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Belemnitella junior belemnite Zone; see Abdel-Gawad, 1986; Walaszczyk et al., 2016). In the
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Maastrichtian type area, it ranges from interval 3 of the Vijlen Member to the upper third of
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the Nekum Member, being associated for almost its entire range with Belemnitella junior
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Nowak, 1913.
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Genus Tenuipteria Stephenson, 1955 (emend. Speden, 1970b)
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Type species. Inoceramus argenteus Conrad, 1858, by original designation.
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Tenuipteria argentea (Conrad, 1858)
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Figs 2A–C, 5E, 6A–H.
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*1858 Inoceramus argenteus Conrad, p. 324, pl. 34, fig. 16.
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*1895 Avicula Geulemensis [sic] Vogel, p. 28, pl. 2, figs 3–5.
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1922 Inoceramus tegulatus v. Hag.; Ødum, p. 1, figs 1–5, 7 (non fig. 6).
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1959 Inoceramus tegulatus v. Hagenow; Dobrov and Pavlova in Moskvin, p. 150, pl. 23, figs
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1979 Tenuipteria geulemensis (F. Vogel, 1895); Dhondt, p. 142 (pars), pl. 1, figs 2, 5–6 (non figs 1, 3–4 = Spyridoceramus tegulatus).
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1–4.
1982 Tenuipteria geulemensis (Vogel, 1895); Dhondt, p. 77 (pars), pl. 1, figs 9–11 (non figs 1–4, ?5–8 = Spyridoceramus tegulatus).
1986 Tenuipteria argentea (Conrad, 1858); Abdel-Gawad, p. 147, pl. 31, figs 3–4 (with
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additional synonymy).
1994 Tenuipteria argentea (Conrad, 1858); MacLeod, p. 1063, fig. 12.1–12.3, 12.7.
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2007 Tenuipteria argentea (Conrad, 1858); Landman et al., p. 35, fig. 12A–G.
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?2007 Inoceramus sp.; Landman et al., p. 35, fig. 14J–M.
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2013 Tenuipteria geulemensis (Vogel, 1895); Gravesen and Jakobsen, p. 64, fig. 1.
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Material. Several dozens of specimens, some articulated, and all preserved in indurated
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biocalcarenites, including NHMM MK 3858, NHMM RZ 02231a, NHMM 2017 004, NHMM
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JJ 13207, NHMM JJ 13408 and Cremers 5576.
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Description. Medium sized, inequivalve and inequilateral. Shell exceedingly thin and sticking
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to matrix when separating internal mould. Left valve inflated, particularly anteriorly; posterior
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wing more flattened. Umbo prosogyrate, slightly incurved; anterior auricle small and
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triangular. Ornament moderately developed, with slightly irregular concentric folds; these are
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intersected by close-set radial furrows, mostly in anterior and median portions of the shell.
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Right valve moderately inflated, umbo projecting slightly above hinge line; auricle small and
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triangular; posterior valve extension broad. Ornament similar to that of left valve, but more
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regular; interspaces between concentric folds and radial furrows smaller. Hinge straight, but
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ACCEPTED MANUSCRIPT ligament mostly either poorly preserved or not exposed. Shell material is mostly greyish pink
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to honey coloured.
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Remarks. When describing Avicula geulemensis, Vogel (1895, p. 29) also noted a ‘... nicht
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sicher bestimmbarer Steinkern dieser Art von Kunraed ....’; this refers very likely to S.
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tegulatus as here understood. In fact, T. argentea does not occur in the ‘Kunrade Limestone’
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(now Kunrade Formation; see Jagt et al., 2017a), but is confined to the higher portion
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(subunits IVf-6 and IVf-7) of the Meerssen Member (Maastricht Formation), with a distinct
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acme in the uppermost 30–50 cm of subunit IVf-6, and ranging upwards into the overlying
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subunit IVf-7, of earliest Danian age (Jagt et al., 2013). Thus, the range is similar to that in
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east-central Poland (Abdel-Gawad, 1986; Walaszczyk et al., 2016), where it is restricted to
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the zone of Belemnella kazimiroviensis and Belemnella skolozdrownae, the first appearance
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datum (FAD) not having been determined yet. However, only in the Maastrichtian type area
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does T. argentea extend into the lowermost Paleocene (compare Ward et al., 1991; MacLeod,
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1994; Marshall and Ward, 1996).
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Occurrence. In addition to earlier records (Dhondt, 1983, 1992; Abdel-Gawad, 1986), T.
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argentea is now also well known from the Gulf and Atlantic coastal plains (Stephenson, 1955;
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Landman et al., 2007) and there is a recent record from the upper Maastrichtian of Jordan
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(Jagt et al., 2017b). The record of Inoceramus sp. by Landman et al. (2007, p. 35, fig. 14J–M)
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appears to be based on a pathological specimen of T. argentea; in material from the
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uppermost Meerssen Member (subunit IVf-6) at Geulhem, a number of individuals show
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more or less comparable growth disruptions.
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4. Discussion As noted by Walaszczyk (in Walaszczyk et al., 2016), the first tegulated inoceramids to appear in the Western Interior of the United States are from the Baculites baculus 10
ACCEPTED MANUSCRIPT ammonite Zone, the base of which equates with the base of the Endocostea typica inoceramid
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Zone. A similar picture is known from east-central Poland, where the earliest members of
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Spyridoceramus are known from Dziurków (Abdel-Gawad, 1986), within the Endocostea
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typica Zone (Walaszczyk, 2004), which correlates with the zone of Belemnella obtusa Schulz,
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1979, Belemnella sp. F and Belemnella sp. G (sensu Remin, 2012).
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As far as the Maastrichtian type area is concerned, the following can be stated. Based on the occurrence of ‘Inoceramus’ ianjonaensis morgani in the upper part of interval 5 and
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the lower part of interval 6 of the Vijlen Member in the Haccourt-Lixhe area (northeast
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Belgium), Walaszczyk et al. (2010, p. 161) noted that this roughly matched the upper
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Spyridoceramus tegulatus/Belemnitella junior Zone of northern Germany (Schulz and
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Schmid, 1983; Schulz et al., 1984; Schönfeld and Burnett, 1991; Schönfeld et al., 1996;
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Christensen et al., 2005). This particular level can also be equated with the Craie à Baculites
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of Manche, northwest France (Walaszczyk and Kennedy, 2011) and the Bay of Biscay
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sections (France, Spain), where ‘I.’ ianjonaensis morgani is known from the lower part of the
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Menuites fresvillensis ammonite Zone (MacLeod, 1994).
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Contrary to Dhondt (1983, fig. 2), T. argentea is here considered to be confined to the upper Meerssen Member, subunits IVf-6 and IVf-7. Such a range would correspond to records
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for her localities ‘Vroenhoven’, ‘Sint Pietersberg’, ‘Geulhem’ and ‘Berg’. However, records
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in the same paper for the localities ‘Eben-Lava’, ‘Lanaye’ and ‘Kunrade’ are here taken to
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refer to S. tegulatus. Stratigraphical ranges, as known to date, of S. tegulatus and T. argentea
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are shown in Figure 7; note that there is no overlap. The ‘flood’ occurrence of T. argentea in
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the uppermost Meerssen Member (subunit IVf-6) is worthy of note and may testify to an
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immigration event. Currently, it cannot be determined whether this event had a westerly (i.e.,
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North America) or an easterly (i.e., Poland or further east) character. In the latter case, it
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might be similar to that documented for the youngest belemnitellid cephalopods (Keutgen et
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al., in press).
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5. Conclusions
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Newly collected material of Spyridoceramus tegulatus and Tenuipteria argentea
allows their stratigraphical ranges to be refined. To date, the former is known to range from
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the Vijlen Member, interval 3 (Gulpen Formation) to at least the uppermost metre of the
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Nekum Member (Maastricht Formation). This range corresponds closely to that in east-central
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Poland (Vistula River valley region), where S. tegulatus occurs in the lower and lower upper
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Maastrichtian (Endocostea typica, Trochoceramus radiosus and ‘Inoceramus’ ianjonaensis
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inoceramid zones, or Belemnella occidentalis and Belemnitella junior cephalopod zones). To
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date, Tenuipteria argentea has only been recorded from subunit IVf-6 of the Meerssen
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Member, with a distinct acme (of both isolated and articulated valves) in the uppermost 20–50
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cm of that subunit. It extends into the overlying subunit IVf-7, which is considered to be of
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earliest Paleocene age (the K/Pg boundary equating with the Berg en Terblijt Horizon), both
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as large-sized articulated specimens in the lower part and as isolated valves (occasionally in
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small clusters) in the topmost 20 cm of the indurated level below the Vroenhoven Horizon
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(Jagt et al., 2013). Except for the extension into the lower Paleocene, this range matches the
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one in east-central Poland where Abdel-Gawad (1986) did not note any overlap in ranges of S.
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tegulatus and T. argentea, a fact corroborated in the present paper. The ‘flood’ occurrence of
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T. argentea in the uppermost Meerssen Member may illustrate an immigration event, but for
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now it cannot be stated if this had a westerly or easterly character. In the latter case, it would
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resemble the migratory patterns of latest Maastrichtian belemnitellid cephalopods of the
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Belemnella kazimiroviensis group.
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Acknowledgements For supply of material, assistance with photography and fieldwork and access to
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quarry grounds, we thank (in alphabetical order): former Ankerpoort (Curfs quarry), CPL SA-
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Haccourt (now Kreco) quarry, Ger Cremers, Mart Deckers, ENCI-HeidelbergCement Group
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quarry (Maastricht), Werner M. Felder(†), René H.B. Fraaije, Loe In de Braekt, Ludo
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Indeherberge, Roland Meuris, Sjir Renkens, Jacques Severijns, John W. Stroucken and Hans
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Zijlstra. Finally, we express our appreciation to the journal reviewers, Ireneusz Walaszczyk
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and Anonymous, for pertinent comments on an earlier typescript.
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the Aachen area (Germany). Netherlands Journal of Geosciences 89, 147–167.
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Captions
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Fig. 1. Map of the extended type area of the Maastrichtian Stage in southern Limburg (the
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Netherlands) and adjacent German (Aachen area) and Belgian (provinces of Liège and
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Limburg) territories, with indication of localities from where either Spyridoceramus tegulatus
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(▲) or Tenuipteria argentea (●) have been recorded to date.
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Fig. 2. Copy (modified) of Vogel’s (1895, pl. 2, figs 3–5) illustrations of Avicula geulemensis;
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A, C – right valve; B – left valve (originals in NCB collections).
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Fig. 3. Spyridoceramus tegulatus (von Hagenow, 1842). A, NHMM 2010 127a (leg. L.
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Indeherberge), CBR-Lixhe quarry, Lixhe (Liège, Belgium); Gulpen Formation, Lanaye
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Haccourt quarry (now Kreco), Haccourt (Liège, Belgium); Gulpen Formation, Lanaye
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Member (flint level 16). D, NHMM JJ 13540 (‘forma C’), CBR-Lixhe quarry, Lixhe (Liège,
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Belgium); Gulpen Formation, Vijlen Member (interval 6), base + 2–4 m. E, NHMM JJ
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15029, ENCI-HeidelbergCement Group quarry, Maastricht, the Netherlands; Maastricht
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Formation, Schiepersberg Member, base + 0.15 m. F, NHMM 2010 127b (leg. L.
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Indeherberge), CBR-Lixhe quarry, Lixhe (Liège, Belgium); Gulpen Formation, Lanaye
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Member (flint level 16). G, NHMM JJ 14803a, ENCI-HeidelbergCement Group quarry,
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Maastricht, the Netherlands; Maastricht Formation, basal Nekum Member. H, NHMM 2017
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003 (leg. R. Meuris), CBR-Lixhe quarry, Lixhe (Liège, Belgium); Gulpen Formation, Lanaye
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Member (flint level 16–21) (see also Fig. 5A). I, NHMM JJ 14238, ENCI-HeidelbergCement
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Group quarry, Maastricht, the Netherlands; Maastricht Formation, basal Nekum Member. J,
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NHMM 2016 048 (leg. R. Meuris), CRB-Lixhe quarry, Lixhe (Liège, Belgium); Gulpen
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Formation, Lanaye Member (flint levels 16–21?), K, NHMM JJ 11965, ENCI-
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HeidelbergCement Group quarry, Maastricht, the Netherlands; Maastricht Formation,
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Gronsveld Member, ENCI Horizon + 0–1.5 m. Scale bars equal 10 mm. All photographs by
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John W. Stroucken, except for D and J which are by Elena Jagt-Yazykova and John Jagt,
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respectively.
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Fig. 4. Spyridoceramus tegulatus (von Hagenow, 1842). A, NHMM 2006 083 (sample W),
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Mosae Forum building pit, Maastricht; Maastricht Formation, Nekum Member, Kanne
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Horizon + 0.3 m. B, NHMM 2017 005 (leg. L. Indeherberge), former CPL SA quarry (now
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Kreco), Haccourt (Liège, Belgium); Gulpen Formation, Lanaye Member (flint levels 17–23).
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C, NHMM JJ 6188a, CBR-Romontbos quarry, Eben Emael (Liège, Belgium); Gulpen
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Formation, Lanaye Member (flint levels 20–21). D–I, copies of Walaszczyk et al. (2010, fig.
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9/2–5, 7–8), all from Gulpen Formation, Vijlen Member, intervals 3, 5 and 6; D–F and H are
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quarry, Lixhe (Liège, Belgium); Gulpen Formation, Lanaye Member (flint levels 17–23). K,
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NHMM JJ 14508, ENCI-HeidelbergCement Group quarry, Maastricht, the Netherlands;
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Maastricht Formation, Valkenburg Member (loose). Scale bars equal 10 mm. All photographs
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by John W. Stroucken, except for D–I and J which are by Ireneusz Walaszczyk and Elena
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Jagt-Yazykova), respectively.
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Fig. 5. Spyridoceramus tegulatus (von Hagenow, 1842) (A–D) and Tenuipteria argentea
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(Conrad, 1858) (E). A, NHMM 2017 003 (leg. R. Meuris), CBR-Lixhe quarry, Lixhe (Liège,
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Belgium); Gulpen Formation, Lanaye Member (flint level 16–21) (see also Fig. 3H). B,
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NHMM JJ 11853 (‘forma C’), CBR-Lixhe quarry, Lixhe (Liège, Belgium); Gulpen
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Formation, Vijlen Member (interval 6), base + c. 6 m. C, NHMM JJ 7011 (‘forma C’), CPL
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SA-Haccourt quarry (now Kreco), Haccourt (Liège, Belgium); Gulpen Formation, Vijlen
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Member (interval 6), base + c. 4 m. D, NHMM JJ 6355, CBR-Romontbos quarry, Eben
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Emael (Liège, Belgium); Maastricht Formation, Nekum Member, middle third. E, NHMM JJ
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13787, former Curfs quarry, Geulhem, the Netherlands; Maastricht Formation, Meerssen
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Member (top of interval IVf-6). Scale bars equal 10 mm. Photographs by John Jagt.
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Fig. 6. Tenuipteria argentea (Conrad, 1858). A–C, NHMM RZ 02231c (ex 663), former
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Curfs quarry, Geulhem, the Netherlands; Maastricht Formation, Meerssen Member (top of
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interval IVf-6). D, NHMM JJ 13408, former Curfs quarry, Geulhem, the Netherlands;
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Maastricht Formation, Meerssen Member (top of interval IVf-7). E, NHMM 2017 004 (ex R.
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Ubaghs Collection), former Curfs (‘Schunck’) quarry, Geulhem, the Netherlands; Maastricht
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Formation, Meerssen Member, interval IVf-6. F, NHMM JJ 13207, former Curfs quarry,
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Geulhem, the Netherlands; Maastricht Formation, Meerssen Member (top of interval IVf-7).
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G, G. Cremers Collection no. 5576, former Curfs quarry, Geulhem, the Netherlands;
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Maastricht Formation, Meerssen Member (top of interval IVf-6). H, NHMM MK 3858 (W.M.
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Meerssen Member (top of interval IVf-6). Scale bars equal 10 mm. All photographs by John
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W. Stroucken, except for D and F which are by Elena Jagt-Yazykova.
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Fig. 7. Regional lithostratigraphy (modified after W.M. Felder, 1996, table 1), with
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indications of stratigraphical ranges of Spyridoceramus tegulatus and Tenuipteria argentea as
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known to date. Note: ‘Vijlen Chalk 0-6’ refers to intervals 0 to 6 of the Vijlen Member.
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S0195-6671(17)30157-X
DOI:
10.1016/j.cretres.2017.05.022
Reference:
YCRES 3619
To appear in:
Cretaceous Research
Received Date: 1 April 2017 Revised Date:
20 May 2017
Accepted Date: 21 May 2017
Please cite this article as: Jagt, J.W.M., Jagt-Yazykova, E.A., Stratigraphical ranges of tegulated inoceramid bivalves in the type area of the Maastrichtian Stage (Belgium, the Netherlands), Cretaceous Research (2017), doi: 10.1016/j.cretres.2017.05.022. This is a PDF file of an unedited manuscript that has been accepted for publication. As a service to our customers we are providing this early version of the manuscript. The manuscript will undergo copyediting, typesetting, and review of the resulting proof before it is published in its final form. Please note that during the production process errors may be discovered which could affect the content, and all legal disclaimers that apply to the journal pertain.
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Stratigraphical ranges of tegulated inoceramid bivalves in the type area of
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the Maastrichtian Stage (Belgium, the Netherlands)
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John W.M. Jagt a,*, Elena A. Jagt-Yazykova b
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a
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b
Uniwersytet Opolski, Katedra Biosystematyki, Pracownia Paleobiologii, ul. Oleska 22, 45-052
Opole, Poland
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Natuurhistorisch Museum Maastricht, De Bosquetplein 6-7, 6211 KJ Maastricht, the Netherlands
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E-mail address: [email protected] (J.W.M. Jagt).
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ABSTRACT
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Recently collected material, in fine- to medium-grained biocalcarenites and in coeval flint
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nodules, of the tegulated inoceramid taxa Spyridoceramus tegulatus (von Hagenow, 1842)
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and Tenuipteria argentea (Conrad, 1858) from various members of the Gulpen Formation and
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the overlying Maastricht Formation allows their stratigraphical ranges to be refined. It is
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concluded that the ranges correspond closely to those in east-central Poland (Vistula [Wisła]
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River valley region), where S. tegulatus occurs in the lower Maastrichtian and lower upper
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Maastrichtian (Endocostea typica, Trochoceramus radiosus and ‘Inoceramus’ ianjonaensis
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inoceramid zones, or Belemnella occidentalis and Belemnitella junior cephalopod zones). In
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contrast, Tenuipteria argentea appears to be confined to the overlying zone of Belemnella
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(Neobelemnella) kazimiroviensis (Skołozdrówna, 1932) and Belemnella (N.) skolozdrownae
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(Kongiel, 1962) and there is no overlap in the range of both species. In the type area of the
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ACCEPTED MANUSCRIPT Maastrichtian Stage, S. tegulatus ranges from the Vijlen Member (interval 3) of the Gulpen
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Formation to the top of the Nekum Member of the Maastricht Formation, while T. argentea is
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restricted to the upper Meerssen Member (subunits IVf-6 and IVf-7, with an acme in the
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former), straddling the Cretaceous-Paleogene (K/Pg) boundary.
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Keywords:
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Mollusca
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Inoceramidae
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Cretaceous
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Stratigraphy
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Correlations
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Northwest Europe
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1. Introduction
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In the extended type area of the Maastrichtian Stage (Fig. 1), inoceramid bivalves generally are rather uncommon and often of mediocre or poor preservation. Notable
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exceptions include assemblages from the Vaals Formation (lower lower Campanian; see Van
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der Weijden, 1943; Sornay, 1982; Walaszczyk and Dhondt, 2005), the Zeven Wegen Member
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(Gulpen Formation; lower upper Campanian) and intervals 3–6 of the Vijlen Member (Gulpen
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Formation, mid- to lower upper Maastrichtian; see Walaszczyk et al., 2010). The youngest
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true inoceramids, ‘Inoceramus’ ianjonaensis morgani Sornay, 1973, Cataceramus
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ghadamesensis (Tröger, in Tröger and Röhlich, 1981) and Endocostea coxi (Reyment, 1955),
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are known from interval 6 of the Vijlen Member, associated with a single tegulated taxon,
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Spyridoceramus tegulatus. Higher upsection, from the Lanaye Member (Gulpen Formation)
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onwards, only thin-shelled, and mostly ill-preserved, tegulated forms remain, occasionally
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occurring in coquina-like concentrations.
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The first tegulated inoceramid to have been recorded from the Maastrichtian type area, although not recognised as such, was Avicula geulemensis Vogel, 1895 (Fig. 2). Of this
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species, now referred to as Tenuipteria argentea, an extensive material has been collected at a
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number of localities in the Dutch and Belgian provinces of Limburg over recent years. The
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same holds true for Spyridoceramus tegulatus, so that the stratigraphical ranges of these taxa
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can now be determined in more detail.
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2. Previous studies
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As documented by Dhondt (1983, 1992), tegulated inoceramids of the genera Spyridoceramus Cox, in Moore, 1969 and Tenuipteria Stephenson, 1955 appear restricted to
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North America (Western Interior, Gulf and Atlantic coasts) and western Eurasia. Dhondt
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(1983) noted that the equivalve S. tegulatus ranged from the lower Maastrichtian to the lower
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upper Maastrichtian, with records from the Western Interior, the European white chalk facies,
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the Russian Platform and central Asia. The inequivalve Tenuipteria argentea was shown to be
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restricted to the upper Maastrichtian, with records from the North American Gulf and Atlantic
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coasts into the European white chalk facies, the Russian Platform and central Asia. Amongst
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named forms, equivalve taxa (see Dhondt, 1983, p. 45) include ‘Inoceramus’ tegulatus,
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Avicula? fibrosa Meek and Hayden, 1856 and ‘I.’ caucasicus Dobrov, 1951, while
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inequivalve taxa comprise ‘Inoceramus’ argenteus, ‘Inoceramus’ dobrovi Jeletzky, in
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Jeletzky and Clemens, 1965 and Avicula geulemensis. This distinction between inequivalve
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and equivalve forms has been referred to in the literature as ‘Avicula-like’ or ‘submytiloid to
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subquadrate’ (i.e., S. tegulatus) and ‘Pholadomya-like’ (i.e., T. argentea). In general,
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ornament in these two taxa is quite similar, although there is a certain range of variation.
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Dhondt (1982, 1983) noted an overlap in the stratigraphical ranges of both taxa in the lower
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part of the Belemnitella junior cephalopod Zone in northern Germany (see also Tröger et al.,
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2001, fig. 13), albeit on the basis of rather poorly preserved material. Abdel-Gawad (1986) noted that there had been some confusion between the two taxa
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amongst previous workers, mostly on account of close similarities in the tile-like (tegulated)
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ornament, taphonomic deformation and the general lack of complete, articulated specimens,
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mainly of S. tegulatus. Unlike Dhondt (1983), Abdel-Gawad did not note any overlap in
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ranges; a notion corroborated in the present paper. In fact, Jagt (1996, 1999) suggested that T.
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argentea was confined to the upper Meerssen Member and that the range of this taxon in the
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Maastrichtian type area corresponded to that in east-central Poland.
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Recently, Walaszczyk (in Walaszczyk et al., 2016, p. 317) has noted that European inoceramid faunas of Campanian and Maastrichtian age were representative of a uniform
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Euramerican Biogeographic Region, and that the final extinction of true inoceramids occurred
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during the early late Maastrichtian (see above). The late Maastrichtian species, referred to as
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tegulated inoceramids (or inoceramoids, sensu Tröger et al., 2001), had invariably been
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regarded to be distinct from genuine inoceramids on features of their ligamental plates.
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However, the loss of regularity in ligament pits had also been seen in a few late Campanian
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and early Maastrichtian inoceramids (Zonova, 1980; Walaszczyk, 2004), so that this specific
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character could not be used for suprageneric classification of such forms (Walaszczyk in
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Walaszczyk et al., 2016).
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These data are here corroborated, and the stratigraphical ranges of S. tegulatus and T.
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argentea in the type area of the Maastrichtian Stage refined. For more details on the
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palaeogeography and lithostratigraphy of the Maastrichtian type area within northwest
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Europe, reference is made to Jagt (1999) and Voigt et al. (2008).
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3. Systematic palaeontology Abbreviations. To denote the repositories of specimens illustrated and/or referred to in the text, the following abbreviations are used: IRScNB, Institut royal des Sciences naturelles
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de Belgique, Brussels, Belgium; NCB, Naturalis Biodiversity Center, Leiden, the
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Netherlands; NHMM, Natuurhistorisch Museum Maastricht, Maastricht, the Netherlands.
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Family Inoceramidae Giebel, 1852
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Genus Spyridoceramus Cox in Moore, 1969 (ex Heinz, 1932, nomen nudum)
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Type species. Inoceramus tegulatus von Hagenow, 1842 (non Ødum, 1922), by original
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designation.
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Spyridoceramus tegulatus (von Hagenow, 1842)
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Figs 3A–K, 4A–K, 5A–D.
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*1842 Inoceramus tegulatus von Hagenow, p. 559.
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*1856 Avicula? fibrosa Meek and Hayden, p. 86.
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*1951 Inoceramus caucasicus Dobrov, p. 167, pl. 2, fig. 2.
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1959 Inoceramus caucasicus Dobrov; Dobrov and Pavlova in Moskvin, p. 150, pl. 19, fig.
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4a–b.
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1970a Tenuipteria fibrosa (Meek & Hayden); Speden, p. 62, pl. 8, figs 1–18; pl. 9, figs 1–16.
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1979 Tenuipteria geulemensis (F. Vogel, 1895); Dhondt, p. 142 (pars), pl. 1, figs 1, 3–4 (non
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figs 2, 5–6 = Tenuipteria argentea). 1982 Tenuipteria geulemensis (Vogel, 1895); Dhondt, p. 77 (pars), pl. 1, figs 1–4, ?5–8 (non figs 9–11 = Tenuipteria argentea).
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1986 Spyridoceramus tegulatus (v. Hagenow, 1842); Abdel-Gawad, p. 146, pl. 31, figs 1–2 (with additional synonymy). 2001 Spyridoceramus cf. tegulatus (Hagenow 1842); Tröger et al., p. 156, text-fig. 12; pl. 1, fig. 6; pl. 3, fig. 2. 2010 Spyridoceramus tegulatus (von Hagenow, 1842); Walaszczyk et al., p. 162, fig. 9/1–8
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(with additional synonymy).
2016 Spyridoceramus tegulatus (v. Hagenow, 1842); Machalski and Malchyk, p. 575, fig. 3D.
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Material. About 50 specimens, preserved both in flint nodules and in fine- to medium-grained
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biocalcarenites, including IRScNB TCMI 10165, IRScNB TCMI 10167, NHMM 2006 083,
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NHMM 2010 127a–b, NHMM 2016 048, NHMM 2017 003, NHMM 2017 005, NHMM
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2017 006a–b, NHMM 2017 007, NHMM JJ 6188a–c, NHMM JJ 6355, NHMM JJ 7011,
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NHMM JJ 11853, NHMM JJ 11965, NHMM JJ 13540, NHMM JJ 14238, NHMM JJ 14508,
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NHMM JJ 14803a–b, NHMM JJ 15029 and NHMM JJ 15895.
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Description. Except for flint-preserved material (e.g., Figs 3A–C, F, H, J, 4B, J, 5A), all
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specimens have suffered from sediment compaction and are fragmentary to varying degrees.
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A number of specimens represent small pieces of shell only, but can be recognised on account
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of their specific ornament. At certain levels, shells occur in clumps, in a coquina-like fashion,
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occasionally as articulated, albeit heavily deformed individuals (e.g., Figs 3K, 4C). For this
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reason, a full description cannot be offered, but specific features of the various shells are
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indicated below.
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Remarks. On the basis of material of S. tegulatus from the Vijlen Member (Gulpen
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Formation) in the Aachen (Germany) and Lixhe-Haccourt (Belgium) areas, Walaszczyk et al.
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(2010, p. 163, figs 2, 9) distinguished three formae, A–C. Their ‘forma A’, occurring in
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interval 3 (Trochoceramus radiosus inoceramid Zone) of that member, is axially elongated
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and slender, with an ornament consisting predominantly of concentric rugae. Radial elements
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concentric rugae intersect. In shell shape and ornament, ‘forma A’ is close to the North
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American Tenuipteria fibrosa, in particular to material from the Fox Hills Formation of South
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Dakota as illustrated by Speden (1970a, b). The lectotype of S. tegulatus from the lower
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Maastrichtian of Rügen (northeast Germany) probably represents the same form (see
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Wolansky, 1933, pl. 5, fig. 6).
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‘Forma B’ is a broader morphotype in which ornament consists nearly exclusively of concentric rugae. This is found in the upper part of interval 5 and in the lower part of interval
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6 (‘Inoceramus’ ianjonaensis inoceramid Zone) of the Vijlen Member, and comes close to
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Inoceramus caucasicus var. ravni Dobrov, 1951. Lastly, confined to the lower part of interval
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6 is ‘forma C’, with very strong radial ornament, in particular anteriorly. This bears a strong
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resemblance to Inoceramus caucasicus var. fistulata Dobrov, 1951 and to one of Wolansky’s
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(1933, pl. 4, fig. 5) syntypes of S. tegulatus.
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be fully assessed in view of the paucity of well-preserved specimens which has so far
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precluded a stratophenetic analysis of the European material. Specimens from the upper
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Lanaye Member (Gulpen Formation) and the Valkenburg, Schiepersberg, Emael and Nekum
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members (Maastricht Formation) generally are of medium size. They all show well-marked
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concentric and radial ornament, covering the anterior and median parts of the shell (where this
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can be determined) or with radial plicae more prominent in the anterior part of the shell.
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Where concentric and radial elements intersect, quadrate to rectangular nodes are seen, best
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developed in the anterior and median parts of the shell. The strong radial ornament (i.e., deep
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grooves anteriorly) of ‘forma C’ is mostly lacking, although a large-sized, flint-preserved left
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valve (Figs 3H, 5A) shows a number of shallow furrows that continue to the ventral margin in
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the anterior portion of the shell. In overall proportions, this individual resembles a specimen
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ACCEPTED MANUSCRIPT of Inoceramus fibrosus from Canada illustrated by Jeletzky (1962, pl. 141, fig. 6), although
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radial plicae are much more prominent, and ornament differs in details. All in all, there is a
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close match between specimens that we have studied and material from the upper
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Maastrichtian of North America (Speden, 1970a, pl. 9, as Tenuipteria fibrosa) and from the
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‘Kunrade Limestone’ (now Kunrade Formation) in the study area as illustrated by Dhondt
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(1979, pl. 1, fig. 3, in particular).
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It would appear that, in shell shape and ornament, the specimens studied by us are uniform (Figs 3A–C, E–K; 4A–C, J–K; 5A, C), although a certain ecophenotypic impact on
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shell morphology may be implied. Where shell (< 1 mm in thickness) material is available,
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the colour is orange brown to dark brown.
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Occurrence. The present species ranges from the lowermost Maastrichtian (upper part of the
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‘Inoceramus’ redbirdensis inoceramid Zone) into the lower upper Maastrichtian (high into the
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Belemnitella junior belemnite Zone; see Abdel-Gawad, 1986; Walaszczyk et al., 2016). In the
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Maastrichtian type area, it ranges from interval 3 of the Vijlen Member to the upper third of
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the Nekum Member, being associated for almost its entire range with Belemnitella junior
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Nowak, 1913.
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Genus Tenuipteria Stephenson, 1955 (emend. Speden, 1970b)
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Type species. Inoceramus argenteus Conrad, 1858, by original designation.
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Tenuipteria argentea (Conrad, 1858)
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Figs 2A–C, 5E, 6A–H.
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*1858 Inoceramus argenteus Conrad, p. 324, pl. 34, fig. 16.
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*1895 Avicula Geulemensis [sic] Vogel, p. 28, pl. 2, figs 3–5.
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1922 Inoceramus tegulatus v. Hag.; Ødum, p. 1, figs 1–5, 7 (non fig. 6).
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1959 Inoceramus tegulatus v. Hagenow; Dobrov and Pavlova in Moskvin, p. 150, pl. 23, figs
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1979 Tenuipteria geulemensis (F. Vogel, 1895); Dhondt, p. 142 (pars), pl. 1, figs 2, 5–6 (non figs 1, 3–4 = Spyridoceramus tegulatus).
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1–4.
1982 Tenuipteria geulemensis (Vogel, 1895); Dhondt, p. 77 (pars), pl. 1, figs 9–11 (non figs 1–4, ?5–8 = Spyridoceramus tegulatus).
1986 Tenuipteria argentea (Conrad, 1858); Abdel-Gawad, p. 147, pl. 31, figs 3–4 (with
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additional synonymy).
1994 Tenuipteria argentea (Conrad, 1858); MacLeod, p. 1063, fig. 12.1–12.3, 12.7.
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2007 Tenuipteria argentea (Conrad, 1858); Landman et al., p. 35, fig. 12A–G.
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?2007 Inoceramus sp.; Landman et al., p. 35, fig. 14J–M.
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2013 Tenuipteria geulemensis (Vogel, 1895); Gravesen and Jakobsen, p. 64, fig. 1.
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Material. Several dozens of specimens, some articulated, and all preserved in indurated
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biocalcarenites, including NHMM MK 3858, NHMM RZ 02231a, NHMM 2017 004, NHMM
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JJ 13207, NHMM JJ 13408 and Cremers 5576.
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Description. Medium sized, inequivalve and inequilateral. Shell exceedingly thin and sticking
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to matrix when separating internal mould. Left valve inflated, particularly anteriorly; posterior
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wing more flattened. Umbo prosogyrate, slightly incurved; anterior auricle small and
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triangular. Ornament moderately developed, with slightly irregular concentric folds; these are
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intersected by close-set radial furrows, mostly in anterior and median portions of the shell.
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Right valve moderately inflated, umbo projecting slightly above hinge line; auricle small and
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triangular; posterior valve extension broad. Ornament similar to that of left valve, but more
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regular; interspaces between concentric folds and radial furrows smaller. Hinge straight, but
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ACCEPTED MANUSCRIPT ligament mostly either poorly preserved or not exposed. Shell material is mostly greyish pink
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to honey coloured.
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Remarks. When describing Avicula geulemensis, Vogel (1895, p. 29) also noted a ‘... nicht
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sicher bestimmbarer Steinkern dieser Art von Kunraed ....’; this refers very likely to S.
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tegulatus as here understood. In fact, T. argentea does not occur in the ‘Kunrade Limestone’
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(now Kunrade Formation; see Jagt et al., 2017a), but is confined to the higher portion
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(subunits IVf-6 and IVf-7) of the Meerssen Member (Maastricht Formation), with a distinct
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acme in the uppermost 30–50 cm of subunit IVf-6, and ranging upwards into the overlying
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subunit IVf-7, of earliest Danian age (Jagt et al., 2013). Thus, the range is similar to that in
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east-central Poland (Abdel-Gawad, 1986; Walaszczyk et al., 2016), where it is restricted to
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the zone of Belemnella kazimiroviensis and Belemnella skolozdrownae, the first appearance
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datum (FAD) not having been determined yet. However, only in the Maastrichtian type area
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does T. argentea extend into the lowermost Paleocene (compare Ward et al., 1991; MacLeod,
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1994; Marshall and Ward, 1996).
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Occurrence. In addition to earlier records (Dhondt, 1983, 1992; Abdel-Gawad, 1986), T.
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argentea is now also well known from the Gulf and Atlantic coastal plains (Stephenson, 1955;
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Landman et al., 2007) and there is a recent record from the upper Maastrichtian of Jordan
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(Jagt et al., 2017b). The record of Inoceramus sp. by Landman et al. (2007, p. 35, fig. 14J–M)
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appears to be based on a pathological specimen of T. argentea; in material from the
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uppermost Meerssen Member (subunit IVf-6) at Geulhem, a number of individuals show
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more or less comparable growth disruptions.
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4. Discussion As noted by Walaszczyk (in Walaszczyk et al., 2016), the first tegulated inoceramids to appear in the Western Interior of the United States are from the Baculites baculus 10
ACCEPTED MANUSCRIPT ammonite Zone, the base of which equates with the base of the Endocostea typica inoceramid
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Zone. A similar picture is known from east-central Poland, where the earliest members of
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Spyridoceramus are known from Dziurków (Abdel-Gawad, 1986), within the Endocostea
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typica Zone (Walaszczyk, 2004), which correlates with the zone of Belemnella obtusa Schulz,
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1979, Belemnella sp. F and Belemnella sp. G (sensu Remin, 2012).
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As far as the Maastrichtian type area is concerned, the following can be stated. Based on the occurrence of ‘Inoceramus’ ianjonaensis morgani in the upper part of interval 5 and
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the lower part of interval 6 of the Vijlen Member in the Haccourt-Lixhe area (northeast
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Belgium), Walaszczyk et al. (2010, p. 161) noted that this roughly matched the upper
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Spyridoceramus tegulatus/Belemnitella junior Zone of northern Germany (Schulz and
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Schmid, 1983; Schulz et al., 1984; Schönfeld and Burnett, 1991; Schönfeld et al., 1996;
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Christensen et al., 2005). This particular level can also be equated with the Craie à Baculites
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of Manche, northwest France (Walaszczyk and Kennedy, 2011) and the Bay of Biscay
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sections (France, Spain), where ‘I.’ ianjonaensis morgani is known from the lower part of the
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Menuites fresvillensis ammonite Zone (MacLeod, 1994).
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Contrary to Dhondt (1983, fig. 2), T. argentea is here considered to be confined to the upper Meerssen Member, subunits IVf-6 and IVf-7. Such a range would correspond to records
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for her localities ‘Vroenhoven’, ‘Sint Pietersberg’, ‘Geulhem’ and ‘Berg’. However, records
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in the same paper for the localities ‘Eben-Lava’, ‘Lanaye’ and ‘Kunrade’ are here taken to
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refer to S. tegulatus. Stratigraphical ranges, as known to date, of S. tegulatus and T. argentea
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are shown in Figure 7; note that there is no overlap. The ‘flood’ occurrence of T. argentea in
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the uppermost Meerssen Member (subunit IVf-6) is worthy of note and may testify to an
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immigration event. Currently, it cannot be determined whether this event had a westerly (i.e.,
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North America) or an easterly (i.e., Poland or further east) character. In the latter case, it
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might be similar to that documented for the youngest belemnitellid cephalopods (Keutgen et
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al., in press).
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5. Conclusions
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Newly collected material of Spyridoceramus tegulatus and Tenuipteria argentea
allows their stratigraphical ranges to be refined. To date, the former is known to range from
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the Vijlen Member, interval 3 (Gulpen Formation) to at least the uppermost metre of the
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Nekum Member (Maastricht Formation). This range corresponds closely to that in east-central
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Poland (Vistula River valley region), where S. tegulatus occurs in the lower and lower upper
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Maastrichtian (Endocostea typica, Trochoceramus radiosus and ‘Inoceramus’ ianjonaensis
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inoceramid zones, or Belemnella occidentalis and Belemnitella junior cephalopod zones). To
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date, Tenuipteria argentea has only been recorded from subunit IVf-6 of the Meerssen
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Member, with a distinct acme (of both isolated and articulated valves) in the uppermost 20–50
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cm of that subunit. It extends into the overlying subunit IVf-7, which is considered to be of
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earliest Paleocene age (the K/Pg boundary equating with the Berg en Terblijt Horizon), both
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as large-sized articulated specimens in the lower part and as isolated valves (occasionally in
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small clusters) in the topmost 20 cm of the indurated level below the Vroenhoven Horizon
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(Jagt et al., 2013). Except for the extension into the lower Paleocene, this range matches the
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one in east-central Poland where Abdel-Gawad (1986) did not note any overlap in ranges of S.
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tegulatus and T. argentea, a fact corroborated in the present paper. The ‘flood’ occurrence of
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T. argentea in the uppermost Meerssen Member may illustrate an immigration event, but for
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now it cannot be stated if this had a westerly or easterly character. In the latter case, it would
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resemble the migratory patterns of latest Maastrichtian belemnitellid cephalopods of the
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Belemnella kazimiroviensis group.
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Acknowledgements For supply of material, assistance with photography and fieldwork and access to
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quarry grounds, we thank (in alphabetical order): former Ankerpoort (Curfs quarry), CPL SA-
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Haccourt (now Kreco) quarry, Ger Cremers, Mart Deckers, ENCI-HeidelbergCement Group
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quarry (Maastricht), Werner M. Felder(†), René H.B. Fraaije, Loe In de Braekt, Ludo
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Indeherberge, Roland Meuris, Sjir Renkens, Jacques Severijns, John W. Stroucken and Hans
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Zijlstra. Finally, we express our appreciation to the journal reviewers, Ireneusz Walaszczyk
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and Anonymous, for pertinent comments on an earlier typescript.
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Captions
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Fig. 1. Map of the extended type area of the Maastrichtian Stage in southern Limburg (the
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Netherlands) and adjacent German (Aachen area) and Belgian (provinces of Liège and
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Limburg) territories, with indication of localities from where either Spyridoceramus tegulatus
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(▲) or Tenuipteria argentea (●) have been recorded to date.
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Fig. 2. Copy (modified) of Vogel’s (1895, pl. 2, figs 3–5) illustrations of Avicula geulemensis;
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A, C – right valve; B – left valve (originals in NCB collections).
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Fig. 3. Spyridoceramus tegulatus (von Hagenow, 1842). A, NHMM 2010 127a (leg. L.
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Indeherberge), CBR-Lixhe quarry, Lixhe (Liège, Belgium); Gulpen Formation, Lanaye
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Haccourt quarry (now Kreco), Haccourt (Liège, Belgium); Gulpen Formation, Lanaye
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Member (flint level 16). D, NHMM JJ 13540 (‘forma C’), CBR-Lixhe quarry, Lixhe (Liège,
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Belgium); Gulpen Formation, Vijlen Member (interval 6), base + 2–4 m. E, NHMM JJ
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15029, ENCI-HeidelbergCement Group quarry, Maastricht, the Netherlands; Maastricht
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Formation, Schiepersberg Member, base + 0.15 m. F, NHMM 2010 127b (leg. L.
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Indeherberge), CBR-Lixhe quarry, Lixhe (Liège, Belgium); Gulpen Formation, Lanaye
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Member (flint level 16). G, NHMM JJ 14803a, ENCI-HeidelbergCement Group quarry,
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Maastricht, the Netherlands; Maastricht Formation, basal Nekum Member. H, NHMM 2017
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003 (leg. R. Meuris), CBR-Lixhe quarry, Lixhe (Liège, Belgium); Gulpen Formation, Lanaye
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Member (flint level 16–21) (see also Fig. 5A). I, NHMM JJ 14238, ENCI-HeidelbergCement
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Group quarry, Maastricht, the Netherlands; Maastricht Formation, basal Nekum Member. J,
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NHMM 2016 048 (leg. R. Meuris), CRB-Lixhe quarry, Lixhe (Liège, Belgium); Gulpen
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Formation, Lanaye Member (flint levels 16–21?), K, NHMM JJ 11965, ENCI-
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HeidelbergCement Group quarry, Maastricht, the Netherlands; Maastricht Formation,
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Gronsveld Member, ENCI Horizon + 0–1.5 m. Scale bars equal 10 mm. All photographs by
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John W. Stroucken, except for D and J which are by Elena Jagt-Yazykova and John Jagt,
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respectively.
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Fig. 4. Spyridoceramus tegulatus (von Hagenow, 1842). A, NHMM 2006 083 (sample W),
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Mosae Forum building pit, Maastricht; Maastricht Formation, Nekum Member, Kanne
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Horizon + 0.3 m. B, NHMM 2017 005 (leg. L. Indeherberge), former CPL SA quarry (now
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Kreco), Haccourt (Liège, Belgium); Gulpen Formation, Lanaye Member (flint levels 17–23).
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C, NHMM JJ 6188a, CBR-Romontbos quarry, Eben Emael (Liège, Belgium); Gulpen
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Formation, Lanaye Member (flint levels 20–21). D–I, copies of Walaszczyk et al. (2010, fig.
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9/2–5, 7–8), all from Gulpen Formation, Vijlen Member, intervals 3, 5 and 6; D–F and H are
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quarry, Lixhe (Liège, Belgium); Gulpen Formation, Lanaye Member (flint levels 17–23). K,
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NHMM JJ 14508, ENCI-HeidelbergCement Group quarry, Maastricht, the Netherlands;
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Maastricht Formation, Valkenburg Member (loose). Scale bars equal 10 mm. All photographs
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by John W. Stroucken, except for D–I and J which are by Ireneusz Walaszczyk and Elena
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Jagt-Yazykova), respectively.
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Fig. 5. Spyridoceramus tegulatus (von Hagenow, 1842) (A–D) and Tenuipteria argentea
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(Conrad, 1858) (E). A, NHMM 2017 003 (leg. R. Meuris), CBR-Lixhe quarry, Lixhe (Liège,
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Belgium); Gulpen Formation, Lanaye Member (flint level 16–21) (see also Fig. 3H). B,
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NHMM JJ 11853 (‘forma C’), CBR-Lixhe quarry, Lixhe (Liège, Belgium); Gulpen
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Formation, Vijlen Member (interval 6), base + c. 6 m. C, NHMM JJ 7011 (‘forma C’), CPL
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SA-Haccourt quarry (now Kreco), Haccourt (Liège, Belgium); Gulpen Formation, Vijlen
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Member (interval 6), base + c. 4 m. D, NHMM JJ 6355, CBR-Romontbos quarry, Eben
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Emael (Liège, Belgium); Maastricht Formation, Nekum Member, middle third. E, NHMM JJ
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13787, former Curfs quarry, Geulhem, the Netherlands; Maastricht Formation, Meerssen
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Member (top of interval IVf-6). Scale bars equal 10 mm. Photographs by John Jagt.
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Fig. 6. Tenuipteria argentea (Conrad, 1858). A–C, NHMM RZ 02231c (ex 663), former
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Curfs quarry, Geulhem, the Netherlands; Maastricht Formation, Meerssen Member (top of
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interval IVf-6). D, NHMM JJ 13408, former Curfs quarry, Geulhem, the Netherlands;
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Maastricht Formation, Meerssen Member (top of interval IVf-7). E, NHMM 2017 004 (ex R.
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Ubaghs Collection), former Curfs (‘Schunck’) quarry, Geulhem, the Netherlands; Maastricht
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Formation, Meerssen Member, interval IVf-6. F, NHMM JJ 13207, former Curfs quarry,
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Geulhem, the Netherlands; Maastricht Formation, Meerssen Member (top of interval IVf-7).
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G, G. Cremers Collection no. 5576, former Curfs quarry, Geulhem, the Netherlands;
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Maastricht Formation, Meerssen Member (top of interval IVf-6). H, NHMM MK 3858 (W.M.
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Meerssen Member (top of interval IVf-6). Scale bars equal 10 mm. All photographs by John
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W. Stroucken, except for D and F which are by Elena Jagt-Yazykova.
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Fig. 7. Regional lithostratigraphy (modified after W.M. Felder, 1996, table 1), with
545
indications of stratigraphical ranges of Spyridoceramus tegulatus and Tenuipteria argentea as
546
known to date. Note: ‘Vijlen Chalk 0-6’ refers to intervals 0 to 6 of the Vijlen Member.
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