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2001 - a Genus Transferred to the Basiliolidae (Pugnacoidea, Rhynchonellida, Brachiopoda)

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2001 - a Genus Transferred to the Basiliolidae (Pugnacoidea, Rhynchonellida, Brachiopoda) BULLETIN DEL’ INSTITUT ROYAL DES SCIENCES NATURELLES DE BELGIQUE SCIENCES DE LA TERRE , 77: 117-129, 2007 BULLETIN VAN HET KONINKLIJK BELGISCH INSTITUUT VOOR NATUURWETENSCHAPPEN AARDWETENSCHAPPEN, 77: 117-129, 2007 Homaletarhynchia SlMON & OWEN, 2001 - a genus transferred to the Basiliolidae (Pugnacoidea, Rhynchonellida, Brachiopoda) by Neda MOTCHUROVA-DEKOVA & Eric SIMON Motchurova-Dekova, N. & Simon, E., 2007 — coquille a contribué également à cette démarche en révélant leur Homaletarhynchia SIMON & O WEN, 2001 - a genus transferred morphologie interne en détail. to the Basiliolidae (Pugnacoidea, Rhynchonellida, Brachiopoda). Bulletin de l'Institut royal des Sciences naturelles de Belgique, Mots-clefs: Crétacé Supérieur, Brachiopoda, Rhynchonellida, Sciences de la Terre, 77: 117-129, 2 pis, 2 figs, Brussels, October Homaletarhynchia, crura subfalciformes, ultrastructure de la 15, 2007 - I S S N 0374-6291. coquille. Abstract Introduction The brachiopod rhynchonellide subgenus Cretirhynchia (Homaletarhynchia) SIMON & OWEN, 2001 is removed from the A first step in the revision of the taxonomically late Cretaceous genus Cretirhynchia P e t t i t t , 1950 and elevated problematic late Cretaceous rhynchonellide brachio­ to a genus level. On the basis o f comprehensive revision o f the pod genus Cretirhynchia P e t t it t , 1950 was made internal morphology o f the type species Terebratulites limbatus von Schlotheim, 1813 by using serial sections, excavations of by S im o n & O w e n in 2001. In their work they the umbonal part and investigations of the shell ultrastructure, made a comprehensive critical review of all the Homaletarhynchia is now placed in the family Basiliolidae, literature dealing with Cretirhynchia. They discussed superfamily Pugnacoidea. For the first time in rhynchonellide exhaustively the various aspects of the “Cretirhynchia taxonomy the peculiarities o f the shell ultrastructure were used to distinguish the representatives of a new genus, removed from problem”: the validity of species and the lack of another one, in which they were originally placed by the founder of knowledge about the internal morphology of many the genus, when the serial sections were not informative enough and species included in the content of Cretirhynchia. before revealing the internal morphology in detail by excavation. These authors published new serial sections of a total for 17 species belonging originally to Cretirhynchia Keywords: Upper Cretaceous, Brachiopoda, Rhynchonellida, Homaletarhynchia, subfalciform crura, shell ultrastructure. and compared them to the serial sections of the type species Rhynchonella plicatilis (J. S o w e r b y , 1816). As a consequence, S im o n & O w e n (2001) subdivided Résumé Cretirhynchia into four subgenera; some of the species were placed in the new genus Woodwardirhynchia, Parmi les brachiopodes rhynchonellides, le sous-genre and some were removed from Cretirhynchia without Cretirhynchia (Homaletarhynchia) SlMON & OWEN, 2001 est retiré du genre Cretirhynchia P e t t i t t , 1950 et est élevé au specifying their new position. All descriptions of the rang de genre. Une révision basée sur la morphologie interne de internal morphologies of the discussed taxa were l’espèce type Terebratulites limbatus VON SCHLOTHEIM, 1813 based on serial sections. The described new genus utilisant des sections transversales sériées, l’excavation de la and four subgenera of Cretirhynchia were said to partie umbonale et l’étude ultrastructurale de la coquille permet be all characterized by raduliform crura, typical of de transférer Homaletarhynchia dans la famille des Basiliolidae au sein de la superfamille des Pugnacoidea. Pour la première the superfamily Hemithiridoidea RZHONSNITSKAIA, fois dans la taxonomie des rhynchonellides, les particularités de 1956. 1’ultrastructure de la coquille ont été utilisées pour distinguer Later M o t c h u r o v a -D e k o v a et al. (2 0 0 7 ) les représentants d’un nouveau genre, lui-même extrait d’un discussed the necessity of using multiple techniques genre plus ancien dans lequel ils étaient placés au préalable par le fondateur du genre, lorsque seules les sections sériées in order to reveal an objective image of the internal n’offraient pas une information suffisante. L’excavation de la characters of some post-Paleozoic rhynchonellides, 118 Neda MOTCHUROVA-DEKOVA & Eric SIMON including representatives of Cretirhynchia. They imaged using a JEOL JSM-6335 F field emission SEM. recommended that a range of methods including The shell thickness and fibres of the secondary layer were optical examination, low vacuum SEM and serial measured close to the symmetry plane. sections should be used whenever possible to fully describe the morphological characters of post- Paleozoic rhynchonellides. They also pointed out that Taxonomy there was a contrast between the working practices of Mesozoic and Cenozoic rhynchonellide workers. Phylum Brachiopoda DUMERIL, 1806 The former prefer serial sections, while the latter, Subphylum RhynchonelliformeaW il l ia m s , working with more poorly consolidated matrix, C a r l s o n , B r u n t o n , H o l m e r & P o p o v , 1996 prefer excavation. These different methods can yield Class Rhynchonellata W il l ia m s , C a r l s o n , contrasting results and a combination of both methods B r u n t o n , H o l m e r & P o p o v , 1996 is preferred in order to describe and illustrate these Order Rhynchonellida KUHN, 1949 taxa more properly. MOTCHUROVA-DEKOVA et al. Superfamily PugnacoideaR zhonsniskaia , 1956 (2 0 0 7 ) first mentioned that subfalciform crura were Family Basiliolidae COOPER, 1959 exposed when investigating some representatives Subfamily Aphelesiinae, COOPER, 1959 of the subgenus Cretirhynchia (Homaletarhynchia). They illustrated the interior of the type specimen Genus Homaletarhynchia SlMON & OWEN, 2001 C. (Homaletarhynchia) limbata (MOTCHUROVA- [Name transferred herein, ex Cretirhynchia DEK.OVA et al., 200 7 ; fig. 5C ), and the difference (Homaletarhynchia) SlMON & OWEN, 2001] between the type species of the nominative subgenus C . (Cretirhynchia,) i.e. C. (Cretirhynchia) plicatilis Diagnosis (with raduliform crura), and the group of species Subpentagonal to subcircular, dorsibiconvex, belonging to C. (Homaletarhynchia) was confirmed symmetrical, medium sized to small rhynchonellides; by shell ultrastructure data. uniplicate, smooth (or very finely striate) with faint or The aim of this paper is to revise the subgenus incipient rounded costae developed only in the anterior Cretirhynchia (Homaletarhynchia) SIMON & OWEN, half. Squama and glotta not developed. Hypothyrid, 2001 with an exhaustive study of its type species umbo short, erect to suberect, foramen small, deltidial Terebratulites limbatus VON SCHLOTHEIM, 1813 plates conjunct, beak ridges distinct, interarea narrow. using all possible methods: serial sectioning, SEM Umbonal part thickened. Convergent to medially observations of dissected umbonal internal structures convex short dental plates, disappearing before the full and investigations of the shell ultrastructure. development of the crura. Strong subquadrate teeth. Euseptoidum well expressed in the dorsal valve. Inner Abbreviations used: IRScNB - Institut royal des socket ridges expanding anteriorly. In juvenile and Sciences naturelles de Belgique, Brussels; NMNHS young individuals crura attached directly to the inner - National Museum of Natural History, Sofia; BMNH socket ridges. Adults develop swollen inner hinge - Natural History Museum, London; L - length of the plates. Crura subfalciform, not curved longitudinally, specimen. distally serrate, short, widening anteriorly as a shovel. Peculiar crater-like negative attachment area developed in the tip of the dorsal umbo. Material and methods Shell ultrastructure Specimens to be macro-photographed were coated with Shell built of two calcite layers primary microgranular ammonium chloride. Serial sections were produced by and secondary fibrous. The fibrous layer is the method summarised by A g e r (1965, pp. 212-218) at a differentiated (not homogeneous), composed of distance of 0.1 mm, subsequently acetate peels were prepared alternating sublayers of two types of fibres: coarser following S t e r n b e r g & B e l d i n g ’s method (1942). and finer. Rhomboidal to subquadrate in cross- Specimens chosen for excavation were manually opened and section coarser fibers prevail. Anisometric anvil­ prepared using steel needle and fine brush. Cross sections at like (=halberd-like) finer fibers are in subordinated the mid shell length for investigation of shell ultrastructure quantity, often developed only close to the primary were polished and etched in 5 % HC1 for 20-30 seconds. Both dissected specimens and the samples for ultrastructure layer. were mounted on stubs then coated in gold palladium and Homaletarhynchia a— basiliolid genus 119 Type species was in fact subfalciform and not raduliform sensu lato Terebratulites limbatus VON SCHLOTHEIM, 1813 as previously presumed. For this reason the diagnosis of the subgenus is here corrected, accordingly. Other species likely included As the kind of the crura determines the placement The list of species which should be included in of this material at a rank above the family level, the Homaletarhynchia will be later discussed elsewhere. whole subgenus must be transferred to another family This list will be based on close similarity in external and superfamily. Therefore, Homaletarhynchia morphology and peculiarities of their serial sections
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