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Multielement Neogondolella ( Conodonta, Upper Permian- Middle

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Multielement Neogondolella ( Conodonta, Upper Permian- Middle Bollettino della Società Paleontologica Italiana Modena, Novembre 1999 Multielement Neogondolella (Conodonta, upper Permian- middle Triassic) Michael J. 0RCHARD Hans RIEBER Geological Survey of Canada Palaomologisches Institut und Museum Vancouver Universitat Zi.irich KEYWORDS- Conodonts, Neogondolella, Multielement apparatus, Permian, Triassic. ABSTRACT- Neogondolella is reconstructed as a multielement conodont genus bearing eight kinds ofelements, seven ofwhich are paired, making a total offifteen elements in al!. The Pa element is segminiplanate, the Pb angulate, Sa is alate, Sbl is breviform digyrate, Sb2 is dolobrate, Scl resembles tertiopedate, Sc2 is bipennate, and M is breviform digyrate. The Sa and Se elements commonly occur as separate anterior and posterior parts. This reconstruction is based on abundant collections ofdisjunct elements and is supported by natura! assemblages ojNeogondolella from Switzerland. The apparatus remained essentially the same from the Upper Permian through the Middle Triassic, with rel4tively minor changes in the morphology o{some elements regarded as ofspecific rank. The apparatus ojNeogondolella diffèrs ftom that ofother Triassic taxa which are nevertheless united in bearing enantiognathiform Sb l elements, on which basis they are classe d as members ofthe supeifamily Gondolellacea. from the Sb elements, the morphology and apparatus pl4n ofgondolel/4ceans generally correspond to that ofthe Ozarkodinida. Within the Jamily Gondolellidae, Gondolella and Neogondo1èlla can be distinguished on multielement grounds. RIASSUNTO- [L apparato di Neogondolella (Con odo ma, Permiano su p.-Triassico medio)]- L'apparato dei conodonti del genere Neogondolella risulta formato da otto diversi tipi di elementi: sette dei quali sono doppi, quindi l'apparato consta di quindici moifoelementi. L'elemento Pa è segminipl4nato, il Pb angulato, l'Sa af4to, Sbl digyrato breviforme, Sb2 do/4brato, Scl sembra terziopedato, Sc2 è bipennato ed M è digyrato breviforme. Gli elementi Sa e Se vengono di norma ritovati con le parti anteriore eposteriore separate. La ricostruzione è basata su abbondanti collezioni di moifoelementi e su associazioni naturali di Neogondolella in Svizzera. L'apparato nel complesso rimane lo stesso dal Permiano superiore al Triassico medio, con alcune modifiche solo nella mor{ologia di alcuni elementi, che vengono considerate di interesse diagnostico a livello specifico. L'apparato di Neogondolella diffèrisce da quelli degli altri taxa Triassici, che hanno l'elemento Sb l enantiognathiforme, e che per questo vengono attribuiti al/4 famiglia Gondoleffacea. Esclusi gli elementi Sb, la moifologia e la struttura dell'apparato dei Gondolelfacea e simile a quel/4 degli Ozarkidinida. Nel/4 jàmiglia Gondolellidae, sul/4 base dell'apparato, èpossibile distinguere Gondolel1a e Neogondolella. INTRODUCTION (Sweet, 1988, fig. 5.28). The common and stratigra- phically useful Triassic elements are almost entirely Triassic conodont apparatuses are relatively poorly assigned to the former family, which includes both known in spite of having been amongst the first to be seximembrate and unimembrate appararuses according interpreted in multielement terms (Huckriede, 1958). to Sweet (1988, p. 87). Other researchers ho l d the view This is due primarily to the extreme rarity ofTriassic that the appararuses of ali gondolellids are multi- natura! assemblage templates, but both morphological element, at least during infancy, and essentially the same convergence and process fragility have rendered in composition (e.g. Kozur, 1989a; Hirsch, 1994a). In statistica! reconstructs difficult. In this paper, the this paper, Neogondolella and other gondolellids are apparatus of the common and long-ranging conodont excluded from the Prioniodinida primarily because Neogondolella is reconstructed. This is accomplished their apparatus does not fearure a digyrate Pb element using collections of well preserved and low diversity but rather an angulate element like the ozarkodinides. discrete elements from Permian and Triassic strata of lt is also recognized that there are severa! different North America and Eurasia. These collections facilitate apparatus types represented amongst the taxa presently an understanding of the gross morphology of the combined within the Gondolellidae. frequently broken ramiform elements, which in rum A diverse generic nomenclarure of gondolellid leads to insights imo how they were configured in the conodonts has developed based on the morphological collapsed apparatuses that are now preserved as features and supposed phyletic relationships of the Pa Neogondolella natura! assemblages in the Middle Triassic platform elements. The proposed Neogondolella of Switzerland (Rieber, 1980; Orchard & Rieber, in multielement apparatus, although relatively conserva- prep.). tive and long-ranging, differs from that of many other A common view ofTriassic conodonts is that most Triassic conodonts and facilitates new ideas on phyletic taxa are closely related variations on a prioniodinide relationships. Multielement reconstructions supporta root stock that diverged in the Late Carboniferous as revised classification, an d strengthen biochronological two families, the Gondolellidae and the Ellisonidae schemes. 476 M.] ORCHARD, H. RIEBER FIRST REPORTS The earliest descriptions of Triassic Neogond- olella-bearing faunas, at that time referred to Gondole/la, were from the Middle Triassic Muschelkalk of Germany (Tatge, 1956). Later, Bender & Stoppe! (1965) introduced the name Neogondolella with N. mombergensis Tatge as the type species (Text-fìg. 1). Concurrent with her description of N mombergensis, Tatge (1956) also described the pectiniform element Ozarkodina ( =Nicoraella) kockeli an d many new ramiform elements, severa! of them incompletely preserved, as single element- or form-species. These ramiform elements were parts ofboth the Neogondolella and Nicoraella apparatuses. The Tatge form-species (Text-fìg.2) that are regarded as parts of the Neo- gondole/la apparatus are listed below: most of these carne from the same stratum and locality. The proposed notation is shown in brackets, with proximal and distai referring to relative position of element parts with respect to the cusp. Text-fìg.1-Neogondolellamombergensis(Tatge, 1956). 1-3) Upper, lower, and latera! views, x80, holotype (Ta 1956/5), a dPa element from the Upper Muschelkalk (1.6 m below top of the Trochitenkalk) in the Schmidtdiel Quarry of Momberg, Marburg District, Germany. THE GENUS NEOGONDOLELLA The genus Gondole/la was introduced for 'gondola- shaped' (segminiplanate) platform elements from the Upper Carboniferous (Stauffer & Plummer, 1932). These elements were characteristically coarsely-ribbed whereas Neogondolella was introduced for the much younger (at the time), morphologically similar bur generally unornamented segminiplanate Permian- Triassic species (Ben der & Stoppe!, 1965). This distinction based on ornament was not universally applicable however, and much subsequent discussion focussed on what criteria could be used to distinguish the two genera (e.g. von Bitter & Merrill, 1977). Although multielement taxonomy featured in these discussions, the uncertainty about gondolellid apparatuses prevented ir from playing a defìning role. Hence, some European workers continue to assign ali species to Gondole/la. Conversely, species formerly Text-fìg. 2 - Neogondolella mombergensis (Tatge, 1956). Sketch referred to Neogondolella have also been subdivided into diagrams of the discrete elements and part-elements numerous additional genera withour reference to named by Tatge and their interpretation as whole multielement composition (e.g. Kozur, 1989b). element components of the Neogondolella apparatus. MULTIELEMENT NEOGONDOLELLA 477 Gondolella mombergensis (P a) recognized that Prioniodella (Angulodus?) prionio- *Ozarkodina tortilis (P b) delloides was the posterior process of Hibbardella *Lonchodina mulleri (M, proximai) *Roundya magnidentata (Sa, proximal) (Roundya) magnidentata, although they did not discuss *Angulodus? prioniodelloides (Sa, distai) the implications of this interpretation (see below). The *Prioniodella decrescens (Sb l, distai) fragmented nature of many ramiform elements also Apatognathus longidentatus (Sbl, proximai) led Kozur & Mostler (1972) to group together Prioniodina insignis (Sb l, proximal) *Metalonchodina mediocris (Sb2, proximal) dissimilar elements, such as those they referred to Metalonchodina digitoformis (M or Sb2, proximai) Hindeodella (Metaprioniodus) suevica: their concept *Prioniodina latidentata (Se, proximai) embraced both the Scl and Sc2 of this paper plus *Prioniodella ctenoides (Se, distai) contiguous Se elements like the holotype of H. suevica, Metalonchodina cf. bidentata (Gunnei) (Sc2, proximal) which is not part of a Neogondolella species. Gen. et sp. indet. C (Scl, proximal) Gen. et sp. indet. B (Scl, proximal) Kozur (197 4, 1976) identified the multielement components of Neogondolella, and indeed gondolellids in generai, as comprising six form-species. These were PREVIOUS MULTIELEMENT RECONSTRUCTIONS (again, with current notation): Huckriede (1958, p. 164) listed eight of Tatge's Gondole/la mombergensis Tatge (Pa) Ozarkodina tortilis Tatge (Pb) elements (shown above with asterisks) under the Enantiognathus ziegleri (Diebel) (Sbl) "conodonten-'Satz"' of Neogondolella navicula, and Prioniodina mulleri (Tatge) (M) thereby introduced the notion of a multielement Metaprioniodus latidentatus (Tatge) (Sc2) Triassic
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