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The Suprageneric Dassifìcation of Some Ordovician Prioniodontid Conodonts

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The Suprageneric Dassifìcation of Some Ordovician Prioniodontid Conodonts Bollettino della Società Paleontologica Italiana Modena, Novembre 1999 The suprageneric dassifìcation of some Ordovician prioniodontid conodonts Svend STOUGE Gabriella BAGNOLI Geologica! Survey Dipartimento Scienze della Terra of Denmark and Greenland Università di Pisa KEYWORDS- Conodonts, Suprageneric taxonomy, Ordovician. ABSTRACT- Phylogenetic relationships among higher taxa within the conodonts that developed a complex apparatus and the resulting classifications are no t universally agreed upon due to the different patterns of the apparatus evolution within the c?ass. Using the most recent reconstructions ofthe prioniodontid apparatuses a picture ofiheir evolution is obtaineCl The proposed classification is base d on diffirent apparatus styles which persisted as unbroken linea_tes. The proposed suprageneric classiftcation for the prioniodontids includes the order Prioniodontida Dzik, 1976 with the supeifamilies Prioniodontoidea Bassler, 1925 and Balognathoidea Hass, 1959. The new order Polyplacognathida with the fomily Polyplacognathidae Bergstrom, 1981 and the new fomily Cahabagnathidae is introduced. RJASS UNTO- [La classificazione sopragenerica di alcuni conodonti prioniodontidi ordoviciani]- Le relazioni filogenetiche e la conseguente classificazione a livello sopragenerico di conodonti con un apparato complesso, sono state oggetto di diffirenti interpretazioni a causa delle (iiverse mod'alità di evoluzione degli apparati nell'ambito della classe. Considerando i più recenti studi sullo stile ed architettura degli apparati dei prioniodontidi, viene qui proposto un diverso modello evolutivo. La classificazione proposta per i prioniodontidi prevede l'ordine .Prioniodontida Dzik, 1976 con le supeifamiglie Prioniodontoidea Bassler, 1925 e Balognath01dea Hass, 1959 ed il nuovo ordme Polyplacognathtda con la fomzglta Polyplacognathzdae Bergstrilm, 1981 e la nuova fomzglta Cahabagnathidae. INTRODUCTION (1988) is used. However, one modifìcation is needed for those pectiniform elements with three main The origin, systematic position, and relationships processes and a secondary postero-lateral process. These between the different groups of the class Conodonta elements are described here as pseudostellate to Pander, 1856 were and still are, controversia!. The distinguish them from pastinate elements with just the recent classifìcations of euconodonts place the genera three primary processes (see Clark et al., 1981, p.W13, with complex apparatuses in the orders Prioniodontida, text-fig.10:1 and text-fig. 10:3). The letter Prioniodinida and Ozarkodinida. The complex abbreviations P, Sand M (Sweet, in Clark et al., 1981; apparatuses com p rise ramiform elements with processes Sweet, 1988) are used in reference to the position of together with coniform elements, blade-like and/or the elements in the apparatus in the same manner as platform-like elements. by Aldrigde et al. (1995). We have not speculated on In this paper, genera with prioniodontid apparatuses the precise orientation of the elements at their respective (Sweet, 1988) and referred to Prioniodontida, are position in the apparatus, because only a few natural reconsidered. The newly gained information dealing assemblages that are considered to be representative of with the biologica! affìnity of the conodonts (Aldridge the Prioniodontida are known (Aldridge et al., 1995; & Donoghue, 1998; Aldridge & Purnell, 1996), the Repetski et al., 1998). function of their skeletal apparatuses (e.g. Aldridge et Some of the representative genera are fìgured. We al., 1993; Aldridge et al., 1995; Dzik, 1991; Nicoll, follow the recommendation 29a of the ICZN (3rd 1987) and elemental growth (Donoghue, 1998) raised edition, 1985) and apply the suffìx -oidea for the a number of taxonomic questions an d new possibilities superfamilies in the systematic section. have arisen. In our treatment we provide diagnoses of the key taxa involved, insofar as they are known and provide HISTORY justifìcation for our taxonomic and evolutionary conclusions. Lindstrom (1970) presented the fìrst modern The descriptive terminology of the elements and suprageneric classifìcation and placed the conodonts the apparatuses given in the Treatise on Invertebrate in the orders Westergaardodinida Lindstrom, 1970 and Paleontology (Sweet, in Clark et al., 1981) and Sweet Conodontophorida Eichenberg, 1930. The two orders 146 S. STOUGE, G. BAGNOLI were further subdivided into eight superfamilies and with genera with the full suite of elements, to the 21 families. Dzik (1976) recognized three suborders Balognathidae Hass, 19 59 as specialized and 14 families. The Treatise (Clark et al., 1981) referred prioniodontides (Lindstrom, 1970; Bergstrom (in the conodonts to the phylum Conodonta Eichenberg, Clark et al., 1981), Sweet, 1988; Aldridge & Smith, 19 30 an d expanded the subdivision of the class 1993; Dzik, 1991). Conodonta Eichenberg, 19 30. In the Treatise New information obtained from work on the Lower classification, the phylum Conodonta and the class to Middle Ordovician conodonts from the Baltoscandic Conodonta were subdivided into two orders, 12 region (Bagnoli et al., 1988; Bagnoli & Stouge, 1997; superfamilies and 48 families. Fahrxus (1983, 1984) Lofgren, 1978, 1985, 1990, 1993a, b, 1994, 1997; criticized the Treatise classification and presented Rasmussen, 1991; Stouge & Bagnoli, 1990; Sturkell, alternative proposals for the phylogeny within the 1991; Zhang, 1997, 1998a, b, c; Zhang & Sturkell, phylum Conodonta Pander, 1856 (Fahrxus, 1983, 1998), Poland (Dzik, 1994; Bednarczyk, 1998), North 1984). America (Stouge & Bagnoli, 1988) and China (Zhang, Sweet (1988) presented a revised version of the 1998a, b) combined with the newly obtained suprageneric classification of the phylum Conodonta information on natura! prioniodontid apparatuses which was different in severa! ways from the older (Aldridge et al., 1995; Aldridge & Theron, 1993; classifications, especially that presented in the Treatise. Gabbott et al., 1995; Repetsk.i et al., 1998), change Sweet (1988) placed the conodonts in a distinct phylum the phylogenetic scenario somewhat and revisions to based on his interpretation of newly-discovered the suprageneric classifications of Sweet (1988), Dzik specimens of the conodont animai (see Aldridge et al., (1991) and Aldridge & Smith (1993) have been 1986; Briggs et al., 1983). In addition, Sweet (1988) proposed (Stouge & Bagnoli, 1998). excluded the protoconodonts and paraconodonts from the phylum Conodonta, and proposed the new classes Conodonti Branson, 1938 and Cavidonti Sweet, 1988. The prioniodontid apparatus Dzik (1991) improved his classification from the one he had presented earlier (Dzik, 1976) and referred The fundamental prioniodontid apparatus is the conodonts to the class Conodonta Pander, 1856 characterized by pectiniform P elements with three with five orders. Of these, the order Prioniodontida primary processes. The apparatus is bilaterally Dzik, 1976 comprised two superfamilies and six symmetrical and sexi- to multimembrate comprising families. two to four kinds of paired P elements, and 9 S Aldridge & Smith (1993) generally followed Sweet elements, one axial and four pairs, and a pair of M (1988) in their outline of the classification of the elements. Advanced prioniodontid apparatuses have Conodonta but also introduced important differences. three or four pairs of P elements. The most complex At that time the knowledge of the anatomy of the prioniodontid apparatus is that of Promissum with four conodont animai had improved (Aldridge et al., 1993) pairs of P elements with two and three primary and information on the zoologica! affinity of the processes (Aldridge et al., 1995). conodont animal pointed towards primitive vertebrates. The prioniodontid apparatus is first recognized in The conodonts were referred to the phylum Chordata Tremadoc strata and radiated in the Ordovician. The Bateson, 1886 and to the class Conodonta Eichenberg, prioniodontid apparatus persisted into the Silurian but 1930 sensu Clark et al., 1981 by Aldridge & Smith declined and disappeared at the end of the Devonian (1993). Aldridge & Donoghue (1998) and Aldridge (Sweet, 1988). & Purnell (1996) presented the most recent overview of the conodonts, and the evidence for the affinity of conodonts with primitive vertebrates related to the Evolution ofPrioniodontida hagfishes has been confirmed (Aldridge & Donoghue, 1998). The oldest representatives with the characteristic prioniodontid apparatus pian are "Acodus"· deltatus Lindstrom, 1955 and Diaphorodus Kennedy, 1980. THE ORDER PRlONIODONTIDA DZIK, 1976 Both taxa are known from the early Tremadoc (Bagnoli et al., 1988; Ethington & Clark, 1981; Kennedy, 1980; The order Prioniodontida Dzik, 197 6 in the recent Lofgren, 1993a; McTavish, 1973; Repetski, 1982; classifications of Sweet (1988), Dzik (1991) and Stouge & Bagnoli, 1988). The two taxa are the first Aldridge & Smith (1993) includes taxa with the representatives of the two major lineages of prioniodontid apparatus composed of elements in P, S Prioniodontida which respectively are reflected in the and M positions. In addition, conodonts forming two superfamilies Prioniodontoidea Bassler, 1925 and bimembrate apparatuses consisting only of pectiniform Balognathoidea Hass, 1959. The youngest Ordovician elements which presumbly occupied P positions were representative of the Prioniodontoidea is probably also included in the Prioniodontida and referred, along Phragmodus Branson & Mehl, 1933, which disappeared ORDOVICIAN
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