pp 061-164 (Jagt-2) 15-01-2007 10:23 Pagina 59 Late Cretaceous-Early Palaeogene echinoderms and the K/T boundary in the southeast Netherlands and northeast Belgium — Part 2: Crinoids John W.M. Jagt Jagt, J.W.M. Late Cretaceous-Early Palaeogene echinoderms and the K/T boundary in the southeast Netherlands and northeast Belgium — Part 2: Crinoids. — Scripta Geol., 116: 59-255, 51 figs., 46 pls, Leiden, May 1999. John W.M. Jagt, Natuurhistorisch Museum Maastricht, Postbus 882, NL-6200 AW Maastricht, The Netherlands, E-mail: [email protected] Key words: Echinodermata, Crinoidea, Late Cretaceous, Early Palaeogene, taxonomy, stratigraphy. All Campanian, Maastrichtian and Danian articulate (cladid) crinoids known to date from the extend- ed type area of the Maastrichtian Stage, are described and illustrated. The geographic and strati- graphic distribution of this unexpectedly diverse echinoderm group are documented. A total of twen- ty genera (three of them new) and thirty-six species (six of them new) are defined: Austinocrinus bico- ronatus (von Hagenow, 1840), ‘Isocrinus’ sp., ‘I.’ lanceolatus (Roemer, 1840)?, Isselicrinus buchii (Roemer, 1840), Praeisselicrinus? limburgicus (Rasmussen, 1961), Nielsenicrinus agassizii (von Hagenow, 1840) (= ‘Pentacrinites’ kloedeni von Hagenow, 1840), N. ewaldi sp. nov., Jaekelometra gr. belgica (Jaekel, 1902), J. gr. concava (Schlüter, 1878) (= J. columnaris Gislén, 1924 and including forma meijeri Rasmussen, 1961), J.? defectiva sp. nov., Placometra gr. laticirra (Carpenter, 1880), Atuatucametra annae gen. et sp. nov., Amphorometra gr. conoidea (Goldfuss, 1839), Semiometra impressa (Carpenter, 1881), S. lenticularis (Schlüter, 1878), S. saskiae sp. nov., Loriolometra retzii (Lundgren, 1875), Hertha gr. pygmea Gislén, 1924, H. gr. plana (Brünnich Nielsen, 1913), H. gr. mystica von Hagenow, 1840 (?), Bourgueticrinus sp. 1 (aff. baculatus Klikushin, 1982c), Bourgueticrinus sp. 2, B. bruennichinielseni Ødum, in Jessen & Ødum, 1923, B. aff. brydonei Rasmussen, 1961, B. constrictus (von Hagenow, in Quenstedt, 1876), B. danicus Brünnich Nielsen, 1913, B. hureae (Valette, 1917), B.? suedicus (Carpenter, 1881), Democrinus? maximus (Brünnich Nielsen, 1915), Monachocrinus? gallicus Rasmussen, 1961, Cyathidium vlieksi Jagt, 1986, Appli- nocrinus cretaceus (Bather, 1924), Birgelenocrinus degraafi gen. et sp. nov., and Veugelersia diana gen. et sp. nov. It is demonstrated that comatulids and bourgueticrinids/bathycrinids in particular show a wide range of variation. Awaiting detailed biometric analyses of modern comatulid populations and a reassessment of comatulid taxonomy, for the time being the use of open nomenclature (Bengtson, 1988) is favoured in these instances. Generic reassignments are proposed for Isocrinus? carinatus Roemer, 1840 [sensu Rasmussen, 1961] and Bourgueticrinus aequalis d’Orbigny, 1841: to the cainocrinid genus Nielsenicrinus and to the otherwise exclusively North American Late Cretaceous genus Dunni- crinus, respectively. Contents Introduction ........................................................................................................................... 60 Material and methods .......................................................................................................... 61 Previous work ....................................................................................................................... 62 Systematic palaeontology .................................................................................................... 67 Acknowledgements ............................................................................................................ 152 References ............................................................................................................................ 153 pp 061-164 (Jagt-2) 15-01-2007 10:23 Pagina 60 60 Jagt. Late Cretaceous and Palaeogene echinoderms, pt 2: Crinoids. Scripta Geol., 116 (1999) Introduction In comparison with other echinoderms, and echinoids in particular, Late Creta- ceous (Campanian-Maastrichtian) and Early Palaeogene (Danian) crinoid faunas from the extended type area of the Maastrichtian Stage have virtually been neglected. Based on a limited number of museum collections only, Rasmussen (1961, 1965) doc- umented a total of sixteen crinoid species from these strata. However, it should be pointed out that he still distinguished four species of the atelecrinid comatulid Jaekelo- metra and that his list includes one species not recorded subsequently from the area, the cainocrinid Nielsenicrinus obsoletus (Brünnich Nielsen, 1913). In the present paper, thirty-six species in twenty genera are described. Three gen- era and six species are new, and a few species are reassigned to other genera. Of note amongst these are in particular new genera and species of pelagic roveacrinids and pterocomid comatulids, as well as cups and large numbers of proximal brachials, which allow Isocrinus? carinatus (sensu Rasmussen, 1961) to be assigned definitely to the cainocrinid Nielsenicrinus. This species may in fact be a member of a phylogenetic lineage leading from the Santonian (-? Early Campanian) ‘Isocrinus’ minutus (Valette, 1917) to the Early Maastrichtian N. agassizii (von Hagenow, 1840) and the Late Maas- trichtian N. rosenkrantzi Rasmussen, 1961. Although collections available for the present paper are large in comparison to previous studies of crinoids from the area, taxonomic assignments are occasionally fraught with difficulties. In these instances, the use of open nomenclature is favoured, awaiting detailed biometric analyses of ‘populations’ of species originally described from elsewhere in northwest Europe. This holds especially true for bourgueticrinids/bathycrinids and some comatulids. In general, a wide range of morphological variation is accepted for these species. For bourgueticrinids in particular a conservative attitude is adopted, although it is real- ised that some taxa (e.g. Monachocrinus? gallicus) may in fact be no more than ontoge- netic stages of other species occurring in the same assemblages. Most of the material available was collected during the past six to eight years in a number of quarries, both working and disused, and in a few other (natural) expo- sures in the area between Aachen, Liège and Maastricht (see Jagt, 1999 for details). Included also are museum collections, despite the fact that these, and especially those made prior to 1975, often suffer from lack of stratigraphic details. A number of pri- vate collections, notably those of M.J. van Birgelen, M.M.M. Kuypers and M.J.M. Deckers, which comprise stratigraphically well-documented material, have also been screened. Specimens from these collections which are illustrated or referred to in the present paper have been transferred to the collections of the Natuurhistorisch Museum Maastricht and have the prefix NHMM. The present paper is the second in a series of contributions documenting echino- derm faunas of Late Cretaceous and Early Palaeogene age from the extended Maas- trichtian type area. For a discussion of the geographic-stratigraphic setting and gener- al references see Part 1 (Jagt, 1999). This, and subsequent papers, are principally taxo- nomic in nature. On the basis of these revisions, the crinoids, echinoids, asteroids, and ophiuroids, will then be analysed palaeobiologically and palaeoecologically in the concluding paper of this series. Analyses of functional morphology in turn will provide explanations for changes in echinoderm faunas across the Cretaceous/Ter- pp 061-164 (Jagt-2) 15-01-2007 10:23 Pagina 61 Jagt. Late Cretaceous and Palaeogene echinoderms, pt 2: Crinoids. Scripta Geol., 116 (1999) 61 tiary (K/T) boundary in the area (Jagt, in prep. d). Comparisons with extant faunas allow conclusions to be drawn with regard to habitat preference and functional mor- phology of these fossil echinoderm taxa. These papers will all comprise a historical account of the research of these echino- derm groups, which is to precede the systematic descriptions, and which will include citations from literature sources which are not easily accessible. Material and methods Over the past six to eight years, members of the ‘Vijlen Groep’, and other (ama- teur) palaeontologists, have taken and processed samples from a number of key sec- tions in the extended type area of the Maastrichtian Stage (see Jagt, 1999). The weight of these bulk samples varied widely and depended mainly on outcrop condition: from a few to more than 500 kg. In their choice of sampling intervals the group members were led primarily by personal observations during fieldwork and by results of bio- clast analyses carried out by P.J. Felder. Spacing of the sampled intervals varied to a certain extent, being largely dependent of accessibility, but coverage was such that this sample series compares well with that described by H. Ernst (1984) for the standard section of the Late Cretaceous white chalk in northwest Germany. Special attention was paid to fossil hash levels which document decreases in sedimentation rates, win- nowing and hard-part concentration. In this way, the entire section from the Early Campanian Vaals Formation to the Middle Danian Geulhem Member was sampled at regular intervals. Stratigraphic provenance of samples was indicated with reference either to the lower or upper boundary level of a given member, depending on outcrop conditions. Invariably, lithostratigraphic logs as published by W.M. Felder (1975b) and W.M. Felder & Bosch (in