Revised Systematics of the Schlumbergerinida (Phylum Foraminifera)
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157 Revised Systematics of the Schlumbergerinida (Phylum Foraminifera) VALERIA MIKHALEVICH1 and MICHAEL A. KAMINSKI2 1. Zoological Institute Russian Academy of Sciences, Universitetskaya nab. 1, St. Petersburg, 199034, Russia. email: [email protected] 2. Department of Earth Sciences, UCL, Gower Street, London, WC1E 6BT, U.K. ABSTRACT __________________________________________________________________________________________ The newly revised systematics of the foraminiferal order Schlumbergerinida is here presented, with diagnoses of all taxa, comprising 21 genera, seven families and eleven subfamilies, of which three fam- ilies (Trilocularenidae, Ammomassilinidae, Ammoflintinidae) and five subfamilies (Psamminopeltinae, Birsteiniollinae, Silicomassilininae, Spirolocammininae, and Pseudoflintininae) are new. A key to the genera and subfamilies of the order is also given. INTRODUCTION the ancestral order to the order Miliolida of the sub- According to new concepts of the foraminiferal class Miliolana. The unity of both orders within a macrosystem and evolution, the former Textulariina single phyletic line is supported by (1) the similari- (sensu Loeblich & Tappan, 1987) represents a het- ty of their morphological features; (2) the profound erogenous group belonging to five different sub- similarity of their ontogenetic and phylogenetic ten- classes: Astrorhizana Saidova, 1981, Ammodiscana dencies; (3) cases of gradual transition from an Mikhalevich, 1980, Miliamminana, Mikhalevich, agglutinated to a porcellaneous secreted wall 1980, Hormosinana Mikhalevich, 1980 and (Mikhalevich et al., 1986, 2004, pl. 1, figs. a, b); (4) Textulariana Mikhalevich, 1980 of the correspon- molecular data (Fahrni et al., 1997, Pawlowski et al., ding classes Astrorhizata Saidova, 1981, Spirillinata 2003). Maslakova, 1990, Miliolata Saidova, 1981, All this makes it necessary to transfer the schlum- Nodosariata Mikhalevich, 1992, and Rotaliata bergerinids from the former Textulariina (or from Mikhalevich, 1980 (Mikhalevich, 1992, 1999, 2000, the order Lituolida, where they were placed later by 2004). The five foraminiferal classes can be charac- Loeblich & Tappan (1992)) to the subclass terised morphologically, and include isomorphic Miliamminana Mikhalevich, 1980 as the separate subclasses with a more primitive shell wall (mostly order Schlumbergerinida Mikhalevich, 1980 agglutinated) and a more advanced one (usually (Jurassic to Holocene). different types of secreted calcareous wall, except Because the representatives of the order among primitive unilocular forms of the class Schlumbergerinida are analogous in their shell and Astrorhizata where the representatives of their chamber form to those of the order Miliolida (as higher subclass Astrorhizana have an agglutinated well as in features of their apertural structures) the wall and those of the lower subclass Lagynana - principles and approaches used in their classifica- have a tectinous wall). Thus, it has been suggested tion ought to be the same to follow classificatory that the transition from a more primitive aggluti- principles at a uniform level. Thus the type of the nated wall to the different types of the fully secret- shell structure (including the mode of coiling, form ed calcareous wall took place independently and in of the chambers and their disposition) is considered parallel lines during geological history, within each as serving as a feature at a family level. The type of class representing a separate phyletic line. The apertural structure (the presence or absence of a majority of agglutinated and calcareous isomorphs tooth, its modification) is used as a feature at the are therefore not the result of convergence as was subfamily level. The same approach was used earli- previously thought, but are in close relationship. er in the classification of the order Miliolida Consequently, isomorphic forms of the subclass (Mikhalevich, 1988). Miliamminana are considered to be the ancestral forms of the subclass Miliolana; the order SYSTEMATICS Schlumbergerinida of the subclass Miliamminana - The new taxonomic position and the composition of In: Kaminski, M.A. & Coccioni, R., (eds), 2008. Proceedings of the Seventh International Workshop on Agglutinated Foraminifera. Grzybowski Foundation Special Publication, 13, 157-166. 158 V. Mikhalevich & M.A. Kaminski the order Schlumbergerinida Mikhalevich, 1980 Family RZEHAKINIDAE Cushman, 1933 designated here is based on the new concepts of Test entirely planispiral; wall agglutinated on an foraminiferal evolution and higher systematics organic base; aperture without a tooth. Cretaceous mentioned above, and on the uniform principles of to Paleocene. their classification for each of the groups, firstly Remarks. Without Rothina, which is considered to within the class Miliolata. The stratigraphic and represent a broken chamber of Caudammina, and is palaeoenvironmental conclusions depend strongly therefore synonymous (Bubík, 1997, Kaminski, upon the taxonomic base. 2004b, Kaminski et al., 1996), and without Ammoflintina, Spirolocammina, Spirosigmoilinella, Subclass MILIAMMINANA Mikhalevich, 1980 Silicosigmoilina, Trilocularena, Silicomassilina placed (= Rzehakinicae Saidova, 1981 = Schlumbergerinana here into other families according to their mode of Mikhalevich, 1992) coiling and shell structure, and Miliammina, Test free, pseudo-twochambered, pseudo-multi- Birsteiniola transferred into Miliammininae. chambered, multichambered, the predominant type of coiling-irregular- and regular glomerate, Subfamily RZEHAKININAE Cushman, 1933 planospiral (often combined with glomerate); shells Test oval and slightly elongated in outline, planispi- of the higher representatives can be cyclical and ral, flattened, but with a thickening at the centre fusulinoid in form; a piece of the tubular chamber because of the partly overlapping chambers, cham- may be preserved around the proloculus; chambers bers half a coil in length, broad, flattened to the predominantly tubular, usually two (more rarely periphery, inflated at their inner side; wall aggluti- three) per whorl, in some forms more wide as in nated on an organic base; aperture without a tooth. Peneroplis-like forms or sometimes subsphaerical, Cretaceous to Paleocene. may be strongly embracing. In higher representa- tives the chamber number greatly increases, cham- Rzehakina Cushman, 1927 ber lumen can be subdivided by the inner skeletal Type species: Silicina epigona Rzehak, 1895 elements, advanced forms may have integrative Test and chambers as described for the family; systems (stolons, tunnels); wall agglutinated on the chambers broad, slightly tapering at the base and at organic base, microgranular or agglutinated and the end, with four to five chambers visible from one partly with secreted needles; aperture terminal, side, only two or two and a half from the other side; simple in the most primitive forms, in more com- wall finely agglutinated; aperture irregularly oval. plex ones - with the outer (flap) or inner tooth of a Cretaceous to Paleocene. special structure, sometimes aperture secondarily multiple, additional apertures non developed; ben- Subfamily PSAMMINOPELTINAE, subfam. nov. thic forms, mostly free living. Nuclear apparatus Test broadly ovoid in outline, flat, fully evolute; and gametes unstudied. Carboniferous to Holocene. chambers half coil in length, narrow, strongly elon- Remarks. The representatives of this subclass often gated, of nearly equal width along the whole length display similarities in form with some of the secret- of the chamber; all chambers visible from both ed Miliolata groups. sides; wall agglutinated on an organic base; aper- ture a subcircular opening. Cretaceous-Miocene. Order SCHLUMBERGERINIDA Mikhalevich, Remarks. The new subfamily differs from the sub- 1980 family Rzehakininae of the same family in being (= ord. Rzehakinida Saidova, 1981) wholly evolute, with non-embracing narrow cham- Test multichambered, usually elongated, of one of bers all visible on both sides of the test. the miliolid types of construction - glomerate at least at the early stage - quinqueloculine, sigmoidal, Psamminopelta Tappan, 1957 (type genus) triloculine, may also be planispiral (mostly spirolo- Type species: Psamminopelta bowsheri Tappan, 1957. culine) or combined, more often with long tubular Test as described for the subfamily; wall finely chambers, two per whorl, more rarely with wider agglutinated; aperture a small subcircular opening. chambers three per whorl resulting in a broader Cretaceous. test; wall agglutinated on an organic base or agglu- tinated and combined with calcareous needles at Subfamily SPIROGLUTININAE Mikhalevich, 1983 the base of the wall; aperture simple or with a tooth Test broadly ovoid in outline, flat, fully evolute; or trematophore. Jurassic to Holocene. chambers half a coil in length, narrow, strongly Remarks. The finding of Miliammina gerochi Tyszka, elongated, of nearly equal width along the whole 1997 in the Middle Jurassic of Poland (Tyszka, 1997) length of the chamber; all chambers visible from extends the known geologic range of this group both sides; wall agglutinated on a calcareous base; from the Cretaceous to the Jurassic. aperture a small circular opening with a short tooth. Miocene to Holocene. Revised Systematics of the Schlumbergerinida 159 Remarks. Placed by Loeblich & Tappan (1987) into Birsteiniolla Mayer, 1974 (type genus) the synonymy of the Hauerinidae, but differs from Type species: Birsteiniolla macrostoma Yankovskaya the representatives