Lanea New Genus, Lineage Ofearly Devonian Conodonts

Bollettino della Società Paleontologica Italiana Modena, Novembre 1999

Lanea new genus, lineage ofEarly Devonian conodonts

Michael A. MuRPHY ]osé Ignacio V ALENZUELA-Rfos Departmenr of Geology Departamento de Geologia Universiry of California a t Davis Universi tar de València

KEYWORDS- Conodonts, Lanea n.gen. , Phylogeny, Early Devonian.

ABSTRACT- Lanea n. gen. is based on a sequence ofPa elements that occurs in the middle Lochkovian centra! Nevada and in the Spanish Pyrenees. In addition, parts of the sequence are known in Alaska, Czech Republic, Austria and Sardinia. The taxa in the lineage enable a subdivision ofmiddle Lochkovian strata that reinforces and refines that suggested by members ojthe genera Ancyrodelloides, Pedavis andFlajsella. The Lanea clade begins at the base ofthe middle Lochkovian with the transjormation ofeosteinhornensis group taxa to L. omoalpha n. sp. (= "Ancyrodelloides omus a." ofMurphy & Matti, 1983). This taxon is interpreted as the progenitor ofL. eoeleanorae n. sp. L. eoeleanorae gives rise to L. eleanorae (Lane & Ormiston, 1979), which in turn gives rise to L. telleri (Schulze, 1968) ( = '1\ncyrodelloides eleanorae "ofKlapper & Murphy, 1980), the latest member of the series. The clade disappears in the above-mentioned regions in the upper part of the trigonicus- pandora Zone towards the end ofthe middle Lochkovian. The transitions within the Pa elements of the lineage show many intermediate morphologies that overlap stratigraphically suggesting an incrementa! mode for the evolutionary changes. During the process the basai cavity, which is open at the onset becomes progressively more restricted. The basai platjorm lobes become terraced and the terraces get progressively larger until they occupy the entire upper surface o{the basai platjorm. In the later jorms, the basai cavity is constricted, the anterior and posterior blades develop benches that extend from the platjorm terrace, and the platjorm terrace develops a rim in the latest described form.

RIASSUNTO- [Lanea n. gen., una linea evolutiva di conodonti del Devoniano inf.] -Il nuovo genere Lanea viene proposto sulla base delle sequenze di elementi Pa ritrovati nel Lochkoviano medio del Nevada centrale e dei Pirenei spagnoli. Parte di questa sequenza è nota anche in Alaska, Repubblica Ceca, Austria e Sardegna. I taxa di questa linea evolutiva consentono di perfezionare la suddivisione Ciel Lochkoviano medio basata sui generi Ancyrodelloides, Pedavis e Flajsella. Il clado Lanea inizia alla base del Lochkoviano medio con la trasformazione di taxa del gruppo eosteinhornensis in L. omoalpha n. sp. ( ='1\ncyrodelloides omus a." Murphy & Matti, 1983). Questo taxon viene considerato il progenitore di L. eoeleanorae n. sp. , da cui discende L. eleanorae (Lane & Ormiston, 1979); quest'ultimo ha dato origine all'ultimo elemnto della sequenza: L. telleri (Schulze, 1968) ( = '1\ncyrodelloides eleanorae " Klapper & Murphy, 1980). Il clado si estingue verso la fine del Lochkoviano medio, nella parte alta della Biozona a trigonicus- pandora La transizione tra i diversi elementi Pa della linea evolutiva mostra varie morfologie intermedie, che si accavallano stratigraficamente, suggerendo una modalità progressiva per le variazioni evolutive. Nel corso del processo, la cavità basale, inizialmente ampia, si riduce progressivamente; la piattaforma diventa terrazzata e queste terrazze si ingrandiscono progressivamente fino a occuparne tutta la parte superiore. Nelle forme tarde la cavità basale è molto piccola, i processi anteriore e posteriore sviluppano delle "panchine '; che si estendono dalle terrazze della piattaforma e, nelle forme più recenti, queste ultime sviluppano un borao rinforzato.

INTRODUCTION the taxon, Ozarkodina n. sp. D [=Ozarkodina delta Klapper & Murphy, 1980; =Kimognathus delta (Klapper This study was initiated to elucidate the early & Murphy) herein], as the zona! name bearer for the phylogeny of Ancyrodelloides, which was represented middle or delta Zone. Valenzuela-Rios & Murphy by Murphy & Matti (1983, fig. 4) as consisting of two (1997, p. 133) suggested that the middle Lochkovian branches, a trigonicus branch and a delta branch, both delta Zone be replaced by a three-fold subdivision of arising out of Ancyrodelloides omus a. Herein, we (l) the interval, the omus a-eleanorae, eleanorae-trigonicus, restrict the genus Ancyrodelloides to the forms that are and trigonicus-pandora Zones. We follow their closely related to A. trigonicus (Tab. l); (2) erect a new classification in this paper (Text-fig. 1), but formalize genus, Lanea, to accommodate the lineage originating the name, omoalpha, for the base of the lowest zone by in Lanea omoalpha n. sp. an d including Lanea eleanorae describing the indicator taxon, L. omoalpha (=omus a (Lane & Ormiston, 1979); (3) erect three new specific- morph Murphy & Matti, 1983). rank taxa, "Ozarkodina"planilingua n. sp., L. omoalpha It should be noted that the lower Lochkovian, the n. sp.(= omus a of Murphy & Matti, 1983), L. interval between the Silurian-Devonian boundary and eoeleanorae n. sp., to accompany Lanea eleanorae and the appearance of L. omoalpha n. sp. is inadequately L. telleri (Schulze, 1968) (=eleanorae of Klapper & zoned. The two taxa used in the Cordilleran region Murphy, 1980) in the new genus Lanea. zonation lack known ancestors or descendents. In Zona! subdivision of the Lochkovian was based on addition, there is an interval between the top of the the scheme introduced by Klapper ( 1977) for the range of "Ozarkodina" eurekaensis and the base of L. Devonian of the Cordilleran region. In i t he employed omoalpha that has not been zoned. 322 M.A. MURPHY, ].I VALENZUELA-RfOS

PRIDOLIAN LOCHKOVIAN PRAGIAN A. murphyi Vaienzuela-Rios, 1994 have been added h e o tra el tri b i s k to this list. The taxa with an asterisk above are assigned to other taxa in this paper. hesperiuseurekaensis eF.F. '.L omoalpha t- The middle Lochkovian strata of centrai Nevada, transitans Oklahoma, Texas, New York, Alaska, the Centrai IN DEX eleanorae 1rigonicus Spanish Pyrenees, centrai Spain (Sierra de SPECIES pandora beta Guadarrama), the Cataionian coastai ranges, Germany gilberti irregularis- (Frankenwaid), Carnic Alps and Karawanken Alps of l praejohnsoni Austr_ia; Bohemia, Czech Republic, Serbia, ltaly delta 1- kindlei (Sardrma) and to some extent, south Australia are 11 11 - 0zarkodina - characterized by the presence of the conodont genus planilingua Ancyrodelloides (Klapper, 1973,1991), which we omoalpha -- interpret as a short-lived offshoot of the spatho- eoele'anorae LANE A gnathodontid eosteinhornensis group of species. The [ l - presence of any member ofAncyrodelloides, wherever i t ltelleri • '- PRIDOLIAN lower middle upper PRAGIAN is found, serves to identifY the middle Lochkovian or basai upper Lochkovian. The middle Lochkovian faunas have become the best known of the Lochkovian Text-fìg. l - Range chart and zonation of the Lochkovian showing the position of the ranges of the species of Lanea new conodont faunas, because of this wide distribution and, genus with respect to taxa used as zona! indices. The consequently, a number of papers have deait with at ranges of the two taxa used as zona! indices for the least a part of the middle Lochkovian fauna (Bischoff delta Zone of the Cordilleran regio n ofNorth America, & Sannemann, 1958; Flajs, 1967; Schulze, 1968; Kimognathus delta and Amydrotaxis praejohnsoni, are also shown far comparison with our zona! scheme. Polsler, 1969; Seddon 1970; Klapper, 1969; Bultynck, Neither of the latter has a known ancestor. 1971; Spassov, 1971; Carls, 1969, 1987; Serpagli et al., 1978; Lane & Ormiston, 1979; Klapper & Johnson, 1980; Klapper & Murphy, 1980; Schonlaub, THE GENUS ANCYRODELLOIDES BISCHOFF & 1980, 1985; Murphy & Matti, 1983; Mastandrea, SANNEMANN, 1958 1985;_ Mawson, 1986; Wang & Zhang, 1988; Sorentmo, 1989; Wilson, 1989; Olivieri & Serpagli, The dose morphological and stratigraphical 1990; Garda-L6pez, et al., 1990; Uyeno, 1990; relationships of Lanea to the genusAncyrodelloides have Vaienzuela-Rios, 1990, 1994; Bardashev & Ziegler, slowed recognition of their separate histories. 1992; Corradini et al., 1998; Ferretti et al., 1998). Ancyrodelloides was originaily set up o n the basis of two Severailineage studies have also been made within the taxa from the Frankenwaid, Germany, A. trigonicus, intervai (Murphy & Cebecioglu, 1986, 1987; Murphy the type species, and A. kutscheri (Bischoff & &._ Springer, 1989). Murphy & Cebecioglu (1987) dealt Sannemann, 1958, p. 91). The two taxa, A. transitans wrth the genus Ancyrodelloides, but di d no t attempt to and A. asymmetricus, described at the same time subdivide the group of morphologies that had (Bischoff & Sannemann, 1958) and regarded by the previously been placed in eleanorae (Lane & Ormiston, originai and subsequent authors (Schulze, 1968; Carls, 1979; Klapper & Murphy, 1980; Murphy & Matti, 1969; Lane & Ormiston, 1979) as closely related toA. 1983). trigonicus, were classified (pre-multielement taxonomy) with the inclusive genus Spathognathodus. With the advent of multielement taxonomy, the taxa in Spathognathodus were transferred to Ozarkodina by default. Murphy & Matti (1983, p. 13) subsequently reclassified these taxa and rediagnosed the genus Ancyrodelloides trigonicus Bischoff & Sannemann, 1958 Ancyrodelloides to include A. transitans, A. asymmetricus, A. transitans (Bischoff & Sannemann, 1958) A. kutscheri Bischoff & Sannemann, 1958 & & A. eleanorae (Lane Ormiston)*, A. delta (Klapper A. asymmetricus (Bischoff & Sannemann, 1958) Murphy)*, A. omus Murphy & Matti, and A. limba- (possibly a variant of A. transitans) carinatus Murphy & Matti* (=Ozarkodina jluminensis A. carlsi (Boersma, 1973) Mastandrea, 1985). Subsequently, Grotsch (1988) has A. omus Murphy & Matti, 1983 included "Spathognathodus"fundamentatus Wailiser* in A. orcula Wilson, 1989 Ancyrodelloides and Wilson (1989) erectedA. orcula for A. secus Barrick & Klapper, 1992 an Australian form that is morphologicaily dose to A. A. cruzae Valenzuela-Rfos, 1994 asymmetricus. Klapper (1991, p. 9) has includedA. carlsi A. murphyi Valenzuela-Rfos, 1994 (Boersma) an d the newly described A. secus Barrick & Klapper (1992). A. cruzei Vaienzuela-Rios, 1994, and T ab. l - List of taxa assigned to Ancyrodelloides. LANEA NEW GENUS 323

THE GENUS LANEA (Murphy & Matti, 1983, pl. 4, fìgs . 20-30). The major faunal elements in the Lochkovian, are lt is seldom possible to give a succinct diagnosis of the simple cones, represented by Panderodus, Belodella, a generic or higher ranking taxon because the later and Pseudooneotodus, the lcriodontidae, the Prionio- members of a clade may bear litde resemblance to their dinidae, and the Spathognathodontidae. The last two ancestors. In the case of Lanea, we regard i t as a separate families have similar apparatuses, bur o n es that are very genus because i t has a separate history. Although, at its different from the other groups. Therefore, there is litde branching point, it is litde different from contem- problem in assigning the apparatus elements to families poraneous taxa, the later forms are distinct. except between the Prioniodinidae and the The characteristic morphology of the derived Spathoghathodontidae. In generai, for those members of the genus Lanea is a robust Pa element apparatuses that are thought to be relatively well with a restricted basai cavity and terraced basai platform established, the prioniodinids have discrete denticles lobe (Text-fìgs. 2, 3). The early taxon, L. omoalpha n. that are aimost circular in cross section, whereas the sp., and closely related variants with an open basai cavity spathognathodontids have denticles that are appressed alllook like members of the eosteinhornensis group of and with more lens-shaped cross sections. However, species, bur they are more robust and the platform lo bes there is some overlap in these and related characteristics. are broadly terraced. They normally differ from For example, it is apparendy a common tendency in Ancyrodelloides in the generailack of ridges or tubercles both families that the deep, open basai cavities and on the basai platform terrace and generaily from the grooves found in the more primitive taxa evolve towards later members ofAncyrodelloides in having lo ba te rather constricted or inverted basai cavities and grooves. The than subquadrate or extensiform laterai processes. The middle Lochkovian genera Ancyrodelloides, Kimogna- genera are similar in that restricted basai grooves and thus, and Lanea follow this generai pattern in the Pa cavities are present in their more derived members. element morphology and, since they commonly occur Lanea differs from the derived members of Kimognathus together, we are stili no t able to reconstruct their entire in lacking a parapet. We include delta in Kimognathus apparatuses. Pb elements have been suggested for Lanea because of the similarity of its Pa element morphology telleri and A. transitans (Lane & Ormiston, 1979; to that of K limbacarinatus (Murphy & Matti) Murphy & Matti, 1983) and these stili seem reasonable.

A 8 c

BASAL GROOVE SECTIONS

Appressed

BASAL Deep Groove VIEW

' 'i,;:ll;:::oove ' lnverted Asymmetrlcal , UPPER VIEW DENTICLE TYPES

---- Fused Discrete' Brim Terrace ' /\/'v\ Triangular DENTI CLE CROSS SECTION JUV/ close •- lens ('(yy' Stout l1 J1 A • ovai equa/ • elliptical /l/VI e clrcular Palisade 11 1/ A /UULwide 4 keeled mm f\/\1\M Needle 11 , J! t1 l r ' r l l l Wl/W/l lrregular Text-fìg. 2 -Nomenclarure of Lanea species. Cross-hatched areas with a left dip represent the pinch zone, those with a right dip represent cross sections of the Pa element. A, B, C are the positions of the cross sections o n the Pa element. 324 MA. MURPHY, ].!. VALENZUELA-RiOS

Sorting out the remaining elements will be diffìcult that normally are assigned to "Ozarkodina" without the discovery of some kind of unusual remscheidensis (Ziegler), but which are probably more occurrence, such as a bedding plane assemblage. closely related to eosteinhornensis or Ozarkodina s.l. because the transition series elements available for reconstruction of the apparatus do no t resemble those HYPOTHESIZED DEVELOPMENT OF THE of remscheidensis (Ziegler, 1960). The sequence of taxa LANEA ELEANORAE LINEACE is based primarily o n the Pa elements of two sections, the Simpson Park Range VII section in Eureka County, The genus Lanea arose out of the plexus of Pa N evada and the Segre l section in the Centra! Spanish element morphologies found in the lower Lochkovian Pyrenees (Text-fìgs. 3, 4). The earliest representatives

SP VII D D • 8993 c

eoeleanorae

B A

omoalpha

Texe-fig. 3 -Simpson Park Range Seceion VII (SP VII). Numbers on ehe left side of ehe column are field locality numbers painted on ehe individuai beds. Numbers o n ehe righe side of ehe column are UCR Museum numbers. The left si de of ehe column is calibraeed in feee, ehe eick marks each represent l O feee; ehe righe side is calibraeed in meeers. LANEA NEW GENUS 325 occur at the base of the middle Lochkovian (as used packstone and contain a variety of shelly foss ils here = base of the omoalpha Zone) of a wide variety of including pelmatozoans, ostracodes, brachiopods, robust Pa elements that are similar in shape and profile tabulate corals, rugose corals, and bryozoans. These to the Pa elements that have been assigned to either beds are normally very lenticular an d are no t traceable remscheidensis or eosteinhornensis by previous authors, laterally even for a hundred meters. The section was but have thickened blades and inflated platform lo bes. trenched. Ali limestone beds in it were sampled for The denticulation is commonly coarser and the cusp conodonts and other shelly fossils and the intervening may be enlarged, but is not appreciably higher than shales were examined for graptolites (Johnson & the other denticles. Pa elements with a small, flattened Murphy, 1969; Berry & Murphy, 1975; Murphy & area next to the biade (= "Ozarkodina" planilingua n. Springer, 1989; Springer & Murphy, 1994). T he lowest sp.; P!. l, figs.l-9) are present in some earlier specimens of Lanea occur just above the uppermost populations as far back as the Late Silurian and are occurrence of Monograptus uniformis Perner and the also common at these levels. In "Ozarkodina" uppermost occurrence of Lanea is just below the lowest planilingua n. sp., the flat area (=terrace; Text-fig. 2) occurrence ofM hercynicus Perner. Essentially the genus was small an d bordered by a narrow br i m (=area Lanea has about the same range as the graptolite M. peripheral to the terrace) of the basai platform. In the praehercynicus Jaeger or approximately the lower two earliest Lanea (=L omoalpha n.sp. 't morph., P!. l, figs thirds of the middle Lochkovian. l 0-19, 23) the area of the terra ce is enlarged. SP-V, the section adjacent to the SP-VII section At higher stratigraphic levels, the terrace is larger and approximately l 00 meters to the south, was used and the angle ofintersection of the brim and the terrace by Klapper ( 1977) to help establish the Lochkovian progressively approached a right angle, the basai cavity is stili open, but the brim is only partially visible in upper view and a granulose surface texture, variably present in earlier morphs, is common on the terrace of the Pa element (=L. eoeleanorae n. sp., P!. l, figs. 20- "'N t l "' 22, 24-26, 30; pl.2, fìgs . 1-11, 21-23). Variants, of the "'o ... N t morph develop higher in the section in Nevada and N l Spain. They (Valenzuela-Rios, 1994; P!. 2, fìgs. 1-6) :l"' l N -;;; • have more centrai platform lobes and fewer, stouter N l ...,o «; . denticles than L. eoeleanorae-r morph. Concurrently, a ·=u · "' c:n ., . "'o. .Q "' in basai sulcus formed in the brim near the lower margin :l E• "' u of the platform. Next, a beneh developed o n the flanks ,._ l l l :.:: 1 :l"' "' ·=o of the anterior and posterior parts of the biade as an Cl) "' Ll.l l "' extension of the platform terrace. At this stage, the brim Ll.l ·E z ... l is no longer visible in upper view and the basai cavity UJ a: is constricted [=L eleanorae (Lane &Ormiston), P!. 2, ?L fìgs. 15-20]. In the next stage, a rim developed at the ..J l outer margin of the terrace and is progressively more f- well defìned. The platform lo bes are constricted at their z "' "o UJ "' "' junction with the biade (especially in mature u c:n l l l o N J: "' :l"' specimens), the basai cavity is markedly constricted, Cl) l l l "' u z

conodont zones of the Cordilleran region. The SP-V Diagnosis- An "Ozarkodina" whose Pa element has and nearby SP-1 sections were studied by Johnson medium-sized basai platform lobes and unconstricted (1973, 1977) who used this areato help establish his basai cavity; open, deep, parallel-sided, anterior basai brachiopod faunal intervals 2 to 4. groove; open, deep, parallel-sided posterior basai groove; The Segre section (Text-fig. 4) in the centrai Spanish area of basai platform terrace small, less than that of Pyrenees is the principal European section used in this brim and meeting brim in obtuse angle; with ascending study and is one of the primary sections used by denticulation posterior of cusp. Valenzuela-Rios (1994) to work out the Lochkovian stratigraphy of the regio n. Like the SP-VII section, the Description- Pa element with mosdy palisade-type Segre section contains allodapic limestone beds, but denticles and smooth surface texture; platform lobes with only a few shale beds. The four taxa in the Lanea with a small terrace extending from biade at various sequence also are present at Segre. The Segre section angles; surface of brim at obtuse angles to terrace an d differs in that L. omoalpha an d L. eoeleanorae enter rh e larger than terrace; basai platform lobes subequal, section together in the lowest sampled bed and there is rounded in upper view and centered slightly more overlap of the four Lanea taxa. posteriorly; basai cavity open; anterior and posterior basai grooves open, deep, parallel-sided; denticles vary in size irregularly with one to three anterior higher; posterior denticles with one or two very small denticles SYSTEMATIC PALAEONTOLOGY then normally three progressively taller denticles before the short denticle at the posterior end; cusp We use the conventional signs and symbols in normally enlarged and high; biade straight or arched synonymy lists as discussed by Matthews, 1973 that in upper view; weak pinch zone present on both were adapted from Richter, 1948. Museum catalogue anrerior and posterior parts of biade. numbers are registered at the U.S. National Museum, Washington (USNM), University of California, Discussion - This taxon is placed provisionally in Riverside (UCR), and Museo Paleont6logico de la Ozarkodina unril the clade characterized by the Universidad de Zaragoza, Spain (MPZ). eosteinhornensis group of species, of which it is probably a member, is formally recognized as a genus. ' Ozarkodina" planilingua n. sp. exhibits a moderate Family SPATHOGNATHODONTIDAE Hass, 1959 range of variation and has been recorded in centrai N evada from the SP-VII, BC-11, and CO P-IV sections Genus 0ZARKODINA Branson & Mehl, 1933 and the centrai Spanish Pyrenees. l t is morphologically most like "Ozarkodina" repetitor Carls & Gandl, an d TJpe species- Ozarkodina typica Branson & Mehl, 1933. "Ozarkodina" eberleini Savage, but has larger basai platform lobes with a small terrace. The genus Ozarkodina is used as the only The specimens figured by Uyeno (1981) resemble convenient existing taxonomic category to house the this taxon in the latera! and basai views given, but it following new taxon until such rime as the relationships can not be discerned in the photographs if they have among members of the eosteinhornensis and the small terrace on the platform lobes, hence the remscheidensis groups of taxa are more completely questionable assignment. worked out and a revised nomenclature is available. T o A more complete synonymy is difficult to give indicate this taxonomic limbo, we have placed beca_use most writers have no t figured ali views of their quotation marks around the generic term Ozarkodina. speCimens.

"0zARKODINA" PLANILINGUA n. sp. Genus LANEA n. gen. P l. l, figs. 1-9 Type species - Ozarkodina eleanorae Lane & ? 19 81 Ozarkodina remscheidensis remscheidensis (Ziegler, 1960) - Ormiston, 1979. UYENO, pl. 3, fìgs. 21-23, 29-33. Diagnosis- Early Pa members of the genus robust Holotype- Specimen UCR 8758, 113 figured on Pl. with medium- to large-sized, terraced, normally l, figs. 7-9 herein, SP-VII Section, Coal Canyon, unornamented basai platform lobes and open basai northern Simpson Park Range, centrai Nevada. grooves, unrestricted basai cavity; later members more robust with appressed or keeled basai grooves, strong Etymology - plani- greek for piane, plus lingua - pinch zone constricted basai cavities, inflated blades tongue - alluding to the small, tongue-shaped terrace and broad, normally unornamented terraces rimmed developed on the basai platform. by sulcate brim. LANEA NEW GENUS 327

Discussion - Lanea is inferred to evolve from the than that of brim and meeting brim in obtuse angle. same root as Ancyrodelloides, on the basis of the Pa and Pb elements, and therefore, the early members Description - Pa element robust with mostly stout, in the two clades have more similarities than the later but some palisade-type denticles and smooth surface members. However, Pa elements of ali taxa we assign texture; platform lo bes shaped like epaulets with terrace to Ancyrodelloides have ridges or tubercles on one or extending from biade a t various angles; surface ofbrim both of the basai platform lobes, whereas only rare at obtuse angles to terrace and narrower than terrace; variants of some of the derived members of Lanea have basai platform lobes asymmetrical, subquadrate- ridges or tubercles. For the later members of the clades, rounded in upper view and centered slightly posteriorly; the processes found in the Pa elements of Lanea are basai cavity open; anterior basai groove open, deep, lobate, whereas those inAncyrodelloides are extensiform. parallel-sided; posterior basai groove open, tapers to a Kimognathus is inferred to be less closely related to point at the posterior end, moderately deep; denticles Lanea an d Ancyrodelloides than they are to each other, vary in size irregularly with o neto three anterior higher although some tendencies, such as, the development and two or three posteri or denti cles enlarged, mid-blade of appressed basai grooves an d constricted basai cavities denticles irregular in size, normally smaller than others; are shared in the derived members of ali three clades. cusp normally enlarged but no t high; upper an d lower In Kimognathus, a parapet paraliel to the biade is profiles slightly concave and slightly convex, developed along one margin of the basai platform of respectively; biade straight or arched in upper view; its derived members. weak pinch zone present on both anterior and posterior The Pb elements that have been suggested an d that parts of biade; normally with smooth terrace surface, are available for reconstructing the apparatuses of ali but specimens known with granulose surface. three genera are robust with lappet-like basai platforms on both inner and outer sides, and anterior and Discussion - Lanea omoalpha exhibits a broad range posterior benches or angulate transverse cross sections of variation and has been recorded in centrai N evada, of the biade (Lane & Ormiston, 1979, pl. l, fig. 47; the centrai Spanish Pyrenees, and the Czech Republic. Murphy & Matti, 1983, pl. 4, figs. 1-3). They resemble The holotype is the model for the tau morph, which is the Pb of Ozarkodina typica more than other the earliest manifestati o n of the generai morphological spathognathodontid Pb elements. They are also similar type. to the Pb elements found in Criteriognathus, but tend A much younger taxon (Pragian) has been identified to have coarser denticles and a higher cusp. They are with the tau morph (= "Ancyrodelloides omus a morph" less similar to the Pb element in the reconstructed of Murphy & Matti, 1983, pl. 2, figs. 18-20) by apparatus of the eosteinhornensis-group and not at ali Mawson & Talent (1994, fig. 11) from the Boola an d like the Pb elements proposed for Amydrotaxis or the Tyers Quarries, Victoria, Australia, but the Australian "Ozarkodina" excavata group of taxa. specimens have longer blades and more even denticulation without the enlarged posterior denticles and apparently without an enlarged cusp, as well as LANEA OMOALPHA n. sp. can be determined from the figures. We do not believe Pl. l, figs 10-19, 23, 27-29; pl. 2, figs. 12-14 that the Australian taxon belongs in Lanea because of the much younger age and the lack of similar morphs 1983 Ancyrodelloides omus alpha morph MURPHY & MATTI, in the intervening strata. We agree with Mawson that p. 17, pl. 2, fìgs. 18-20 [imprint 1982]. her taxon is very dose to L. omoalpha morphologically, 1994 Ancyrodelloides omus morphorype oV AL ENZUEIA-Rfos, pl. l, fig. 10. but believe that it is a heterochronously convergent non 1994 Ancyrodelloides omus al p ha morph, Murphy & Matti, form. 1983- MAwsoN & TALENT, p. 51, fìgs. llH-L. L. omoalpha normally has a smooth terrace, but specimens are known from Spain that have a granulose Holotype- Specimen UCR 8762, 1/1, p. 5, figured surface on the terrace. o n p l. l, figs. 17-19 herein, SP VII Section, Co al Canyon, northern Simpson Park Range, Nevada. LANEA EOELEANORAE n. sp Etymology- omo- greek for shoulder, plus alpha P l. l, figs. 20-22, 24-26, 30; pl. 2, figs. 1-11, 21-23 - beginning letter of the greek alphabet, alluding to the beginning member of a shouldered series of taxa. 1992 Cruciodus eleanorae (Lane & Ormiston, 1979) - BARDASHEV & ZI EG LER, pJ. 2, fìgs. 4, 5, 10. 1980 Ozarkodina cf. asymmetrica Bischoff & Sannemann- Diagnosis - A Lanea whose Pa element has large ScHONIAUB, p!. 4, fìgs. 3, 4. basai platform lobes and unconstricted basai cavity; open, deep anterior basai groove; open, moderately Holotype - Specimen UCR - 9060, 1/2, SP VII deep posterior basai groove tapering uniformly to a Section, Coal Canyon, northern Simpson Park Range, point, area ofbasal platform terrace equal to or greater Nevada, figured on Pl. l, figs. 20-22, 30. 328 M.A. MURPHY, J! VALENZUELA-RfOS

Diagnosis - A Lanea whose Pa element has a distinct brim sulcus and nearly right angle junction platform with a terrace extending at nearly right angles of the brim and terrace, with a pinched biade, but from the biade and meeting the brim at nearly right the basal grooves are not closed. angles, strongly pinched along the base of the element, lt is known from Nevada and Spain. but with open basal cavity and basai grooves; posterior basal groove tapers to a point posteriorly, anterior basal groove parallel sided; biade without bench. LANEA ELEANORAE Lane & Ormiston, 1979 P l. 2, fìgs. 15-20 Description - Pa element robust commonly with granulose surface texture on the terrace; terraces ofbasal 1979 Ozarkodina eleanorae lANE & 0RMISTON, p. 55, pl. l , fìg. 40, not fìg. 47; pl. 2 fìgs. 6-7; pl. 3, figs. 7-8, 11- platform lo bes meet surface of brim an d biade nearly 12. at right angles; outer margin of terrace abruptly non 1980 Ozarkodina eleanorae Lane & Ormiston - l

EXPLANATION OF PLATE l

Ali figures X33. Convention for UCR catalogue numbers is Section followed by Bed Number followed by siide and specimen number; for MPZ the convention is Section Number followed by bed number.

Figs. 1-9 - "Ozarkodina" planilingua n.sp. 1-6) Birch Creek-II Section, UCR 943, Il l , I/2, northern Roberts Mountains Nevada; 7-9) hoiorype, t morph, SP-VII Section, UCR 8758, I/3, Coai Canyon, centrai Nevada.

Figs. 10-19, 23, 27-29 - Lanea omoalpha n.sp. lO) Simpson Park VII Section, UCR 8762, X/7; 11-12) UCR 8762, X/l; 13-15, UCR 8762, I/4; 16, 23) UCR 7912, I/10; 17-19) hoiorype, UCR 8762, I/11, Coai Canyon, centrai Nevada; 27 -29) MPZ 8600, Gerri l.l/17E, centrai Spanish Pyrenees, Spain.

Figs. 20-22, 24-26, 30 - Lanea eoeleanorae n.sp. 20-22, 30) hoiorype, Simpson Park VII Section, UCR 9060, I/2; 24-26) a morph, UCR 9391, I/30, Coai Canyon, centrai Nevada. MA. MURPHY, JI. VALENZUELA-RfOS, LANEA NEW GENUS P!. l 330 M.A. MURPHY, ]I VALENZUELA-RiOS anterior and posterior basai grooves, restricted basai The specimen figured by Bardashev & Ziegler cavity; rimless, terraced, basai platform lobes, (1992) is excluded because it lacks benches on either semicircular in upper view that extend as benches aiong the anterior or posterior biade and the brim is visible the anterior and posterior blades. in oral view. L. eleanorae is a widely distributed, although not Description - Pa element robust with granulose especiaily abundant taxon in the middle Lochkovian. surface texture and platform lo bes shaped like epaulets; lt has been identified in Nevada, Spain, Austria and platform terrace flat, meets brim at abruptly rounded Tajikistan in addition to its type area in Alaska. shoulder, brim pinched or constricted under terrace so that a sulcate area follows margin ofbasai platform lo be and constricts basai cavity; basai platform lo be in upper UNEA TELLERI (Schulze, 1968) view semicircular, constricted or not at junction with Pl. 2, figs 24-40 the biade and centraily located o n biade; terrace extends aiong both anterior an d posteri or blades to form bench v* 1968 Spathognathodus steinhornensis tel!eri ScHULZE, p. 229, pll7, fìgs. 18-19. at base of denticles; denticles mostly discrete except 1978 Spathognathodus steinhornensis te/Ieri Schulze- SERPAGLI, near their bases, stout, may be peg-like and nearly GNou, MAsTANDREA, & OuVIERI, p. 308, pl. 27, fìgs. circular in cross section; biade straight, slightly bent 2, 6. posteriorly, or slightly sigmoidai in upper view; basai 1980 Ozarkodina eleanorae Lane & Ormisron - KiAPPER & grooves parailel-sided, appressed. MURPHY, fig. 4, no. l, 7-9, 13, 14. 1980 Pandorinellina steinhornensis telleri (Schulze) - ScHONLAUB, pl. 2, fig. 20. Discussion-The presence ofbenches aiong the biade 1983 Ancyrodelloides eleanorae (Lane & Ormisron) - MuRPHY with appressed basai groove and constricted basai cavity & MATTI, pl. 4, fìgs. 1-6. separate L. eleanorae from its progenitor, L. eoeleanorae; 1985 Ozarkodina steinhornensis tel!eri Schulze - MASTANDREA, p!. 4, fìgs. 2-4, 6. the lack of a rim at the platform shoulder and centrai 1988 AmydrotttXis johnsoni (Klapper, 1969) aipha morphotype position of the platform lobes and basai cavity - WANG & ZHANG, p. 150, pl. l, fìgs. 9, 10. distinguish the taxon from L. telleri (Schulze). 1991 Ancyrodelloides eleanorae (Lane & Ormisron, 1979) - The width of the ledges and extension posteriorly KiAPPER, Ancyrodelloides, 2, fìgs. 3a,b. of the platform terrace show significant variation in 1994 Ancyrodelloides eleanorae (Lane & Ormisron) - V ALEN- ZUELA-Rfos, p.50, p!. 4, fìgs. 10-12, 14-16. the specimens figured by Lane and Ormiston, (1979). 1998 Pandorinellina steinhornensis tel!eri (Schulze) - CoRRA- Lane & Ormiston (1979, pl. l, fig. 47) assigned a DINI, FERRETTI, & 5ERPAGLI, p. 170, pJ. 2.3.1, fìg 6. Pb element to L. eleanorae that, following Murphy & Matti (1983, p. 23), we believe is probably part of Holotype- Specimen figured by Schulze, 1968, the A. transitans apparatus. p l. 17, figs. 18-19, deposited a t the Geologica!-

EXPLANATION OF PLATE 2 Ali fìgures X 33.

Figs. 1-11, 21-23 - Lanea eoeleanorae n. sp. 1-3) a morph, Simpson Park Range Section VII, UCR 9391, III/34, Coai Canyon, centrai Nevada; 4-6) MPZ 8002, Segre 117; 7) morph, MPZ 8601, Segre 113, centrai Spanish Pyrenees; 8-10) 't morph, Simpson Park Range VII, UCR 8767, I/50; 11) UCR 9393 II/l, 21-23, UCR 8767 I/50, Coai Canyon, centrai Nevada.

Figs. 12-14 - Lanea omoalpha n. sp. 12-14) MPZ 8006, Segre 113, centrai Spanish Pyrenees.

Figs. 15-20 - Lanea eleanorae (Lane & Ormiston) 15-16) Simpson Park Range Section VII, UCR 9393 I/4, bed llB; 17) bed llL I/25, Coai Canyon, centrai Nevada; 18-20) MPZ 8227, Segre 116, centrai Spanish Pyrenees.

Figs. 24-40 - Lane a te/Ieri (Schulze, 1968). 24-25) Simpson Park Range Section VII, UCR 9417, III/4, III/l; 26-27) UCR 8993, III/17, III/16; 28-30) UCR 9417, III/3; 31-33) UCR 9397 V/44; 34-35) UCR 9397, V/45, Coai Canyon, centrai Nevada; 36-38) Gerri de la Sai,l.l/20, MPZ 8060, centrai Spanish Pyrenees; 39,-40) Pb elements, Simpson Park Range Secrion VII, UCR 9397 MA. MURPHY, ].I. VALENZUELA-RfOS, LANEA NEW GENUS P!. 2 332 M.A. MURPHY, JI. VALENZUELA-RiOS

Palaeontological lnstitute, University of Ti.ibingen, N evada have narrow, elongate, rimmed platform lo bes Germany. that extend out perpendicularly from the biade. These are connected by intermediate sized specimens wirh Revised Diagnosis- A Lanea whose Pa element has intermediate morphologies to the maximum-sized a constricted basai cavity, platform terrace with a rim specimens and so are interpreted to be growth stages and anterior and posterior benches on the biade; basai of the same taxon (Pl. 2, fìgs. 24-38). The juvenile cavity posterior of center. holorype from the Karawankian Alps resembles juveniles from Nevada in denticulation and narrow, Description - Pa element robust, with arched or elongate plarforms, but its platform has no marginai straight blade, may be slightly ben t posteriorly; smooth rim and rhe platform lobes make an acute angle with surface texture; asymmetrical subequal platform lobes rhe biade rarher rhan a right angle. aimost entirely posteri or of midpoint ofblade; platform surface rimmed by narrow marginai ridge situated at Distribution - L. telleri has been found in the edge of platform in upper view; ridge extends along Karawankian and CarnicAlps, Sardinia, China, Spain, margins of anterior and posterior ledges; basai cavity and Nevada. Generally, it is associated with the typical dumb-bell shaped and constricted to about haif the middle Lochkovian fauna of the eleanorae-trigonicus size of the upper platform surface in mature specimens; Zone, Flajsella species, Ancyrodelloides transitans and anterior and posteri or grooves, parailel-sided, appressed; A. trigonicus. 13-17 almost uniform mostly palisade-type denticles, with anterior ones higher, descending incrementally to cusp, cusp may or may not be enlarged; anterior and posterior rwo denticles may be somewhat larger. ACKNOWLEDGEMENTS Pb element robust, angulate pectiniform element slightly rwisted; like Pa element, palisade denticles W e rhank Perer Carls and Rurh Mawson for careful reviews of appressed, platform extends subequaily on inner and the paper. Murphy's work was supporred by rhe Narional Science outer sides like lappets, and with bench on both Foundarion and rhe Universriy of California Imramural Research Fund; Valenzuela-Rfos' work was supporred by the Fulbright processes. Foundarion an d rhe Alexander von Humboldr-Srifrung. Discussion- L. telleri was formerly classifìed by some authors with L. eleanorae (Klapper & Murphy, 1980; Murphy & Matti, 1983; Murphy & Cebecioglu, 1987; Klapper, 1991), but the rwo taxa have only a smaii REFERENCES overlap in stratigraphic range andare morphologically BARDASHEV, I. A. & ZIEGLER, W., 1992, Con odo m biosrrarigraphy distinct. L. telleri differs from L. eleanorae in the posi ti o n ofLower Devonian deposits of rhe Shishkat Secrion (Sourhern of the basai platform lo bes and basai cavity, which are Tien-Shan, Middle Asia): Courier Forschungsinstitut more posterior; the pinched shape of the platform lo bes Senckenberg, 154: 1-30, 6 pls. where they join the biade (in mature specimens); the BARRICK,]. & KlAPPER, G., 1992, Late Silurian and Early Devonian conodonrs from rhe Hunron Group (Upper Henryhouse, smooth surface texture of the Pa element, which is Haragan, and Bois d'Are Formations), Sourh-Cenrral granulose in L. eleanorae and the basai platform lobes Oklahoma. In Barrick, J.E. & Chapin, J (eds.), Special Papers of L. telleri have a rim. in Paleomology and Stratigraphy: A Tribute to Thomas W. The holotype of L. telleri is a smaii juvenile with a Amsden: Oklahoma Geologica! Survey, Bulletin, 145: 19-65, 6 pls. morphology that is slightly different from the majority BERRY, W B. N. & MURPHY, M. A., 1975, Silurian and Devonian of rhe juvenile specimens assigned to rhe species. lts graprolires of centrai Nevada: University of California stratigraphic position an d generai characreristics suggest Publications Geologica! Sciences, 110: 109 p., 15 pls. that i t is an extreme variant with respect to the angle at BISCHOFF, G. & SANNEMAN N, D., 1958, Unrerdevonische which the platform lobes intersect the biade and that Conodomen aus dem Frankenwald: Notizblarr hessischen Landes-amre fii.r Bodenforschung, 86: 87-110, 4 pls. the lack of a marginai rim berween the brim and the BuLTYNCK, P., 1971, Le silurien supérieur et le Dévonien inférieur terrace in the holotype may be because the rim has not de la Sierra de Guadarrama (Espagne cenrrale). Deuxième yet developed in rhe very early ontogenetic stage that parti e: assemblages de Conodomes à Spathognathodus: Institut the specimen represents. Only rwo other juveniles have royal des Sciences naturelles de Belgie, Bullerin, 47: 1-43, 11 pls. been figured that are like Schulze's holotype CARLS, P., 1969, Die Conodonren des tieferen Unrerdevons der (Schonlaub, 1980, pl. 2, fìg. 20; Mastandrea, 1985; Guadarrama (Mirrel-Spanien) und die Srellung des pl. 4, fìg. 5). Grenzbereiches Lochkovium/Pragium nach der rheinischen Juveniles of rhis taxon are common in many sections Gliederung: Senckenbergiana Lerhaea, 50: 303-355, 4 pls. (e.g. CarnicAlps) and may become an important factor CARLS, P., 1987, Ein Vorschlag zur biostratigraphischen Redefinition der Grenze Gedinnium/Siegenium und in correlaring fìne grained rocks of mid Lochkovian benachbarrer Umer-Srufen. Strarigraphische Argumeme und age. The juveniles associated wirh fully developed Korrelationen: Courier Forschungsinsrirut Senckenberg, 92: 77- specimens in SP VII and Mill Canyon sections of 121. LANEA NEW GENUS 333

CoRRADINI, C., FERRETTI, A. & SERPAGLI, E., 1998, An Early MuRPHY, M.A. & CEBECIOGLU, M.K., 1986, Statistica! srudy of Devonian section near Fluminimaggiore (Galemmu). In E. Ozarkodina excavata (Branson & M ehi) and O. tuma Murphy Serpagli (Ed.) Sardinia Field Trip Guidebook, Seventh & Matti (Lower Devonian, delta Zone, conodonts, Nevada): International Conodont Symposium h el d in Europe: 168-174, Journal of Paleontology, 60 (4): 865-869. l p!., MuRPHY, M .A. & CEBECIOGLU, M.K., 1987, Morphomerric srudy FERRETTI, A., CoRRADINI, C. & SERPAGLI, E., 1998, The Silurian- of the genus Ancyrodelloides (Lower Devonian conodonts) Devonian Sequence in SW Sardinia. In E. Serpagli (Ed.) centrai Nevada: Journal ofPaleontology, 61: 583-594. Sardinia Field Trip Guidebook, Seventh International MuRPHY, M.A. & MATTI, ].C., 1983, Lower Devonian conodonts Conodont Symposium held in Europe: 57-61. 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Windmill Limestone of centrai Nevada: Journal of RicHTER, R., 1948, Einfiihrung indie Zoologische Nomenklarur Paleontology, 47: 1013-1030, 5 pls. durch Erlauterung der Internationalen Regeln. 2 Aufl.: 1-252, J OHNSON, J. G., 1977, Lower an d Middle Devonian fauna! intervals Frankfurr am Main. in centrai Nevada, based on brachiopods. In Murphy, M.A., ScHONLAUB, H.-P., 1980, Field Trip A, Carnic Alps, Guidebook Berry, W.B.N., & Sandberg, C.A. (eds.) Western Norrh & Abstracts, Second European Conodont Symposium, ECOS America: Devonian. University of California, Riverside II: 5-60, 25 pls. Campus Museum Contribution, 4: 16-32. ScHONLAUB, H.-P., 1985, Devonian conodonts from the section ]OHNSON,].G. & MuRPHY, M.A., 1969, Age and position ofLower Oberbuchach II in the Carnic Alps (Austria): Courier Devonian graptolite zones relative to the Appalachian standard Forschungsinstitut Senckenberg, 75: 353-374, 6 pls. succession: Geologica! Sociery of America, Bulletin, 80: 1275- ScHULZE, R., 1968, Die Conodonten aus dem Palaozoikum der 1282. mittleren Karawanken (Seeberggebiet): Neues Jahrbuch KLArrER, G., 1969, Lower Devonian sequence Royal Creek, Yukon Geologie-Palaontologie, Abhandlungen, 130 (2): 133-245, Territory and Devo n Island, Canada with a section o n Devo n 20 pls. Island stratigraphy by A. R. Ormiston: Journal ofPaleontology, SEDDON, G., 1970, Pre-Chappel conodonts of the Llano region, 43: 1-27, 6 pls. Texas: Bureau ofEconomic Geology, Reporr oflnvestigations, KLAPPER, G., 1973, Ozarkodina. In Ziegler, W (ed.), Catalogue of 68: 1-130, 19 pls. Conodonts, Volume l: E. Schweizerbarr'sche Verlagsbuch- SERPAGLI, E., GNOLI, M., MASTANDREA, A. & OUVIERI, R., 1978, handlung, Srurrgart: 228, 249, 2 pls. Paleontological evidence of the Gedinnian (Lower Devonian) KLAPPER, G., 1977, Lower and Middle Devonian sequence in in southwestern Sardinia: Rivista Italiana di Paleontologia e centrai N evada. 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I., 1990, Lochkovian conodonts an d MASTANDREA, A., 1985, Early Devonian (Lochkovian) conodonts srratigraphy a t Gerri de la Sal (Pyrenees): Courier Forschungs- from southwestern Sardinia: Bollettino della Società institut Senckenberg, 118: 53-63. Paleontologica Italiana, 23: 240-258, 5 pls. VALENZUELA-Rfos, J. L, 1994, Conodontos del Lochkoviense y MATTHEWS, S.C., 1973, Notes on open nomenclarure and on Praguiense (Dev6nico Inferior) del Pirineo centrai espafiol: synonymy lists: Palaeontology, 16 (4): 713-719. Memorias del Museo Palaeont6logico de la Universidad de MAwsoN, R., 1986, Early Devonian (Lochkovian) conodont faunas Zaragoza, 5: 1-192, 9 pls. from Windellama, New South Wales: Geologica et V ALENZUELA-Rfos, J. L & MuRPHY, M. A., 1997, A new zonation Palaeontologica, 20: 39-71. of middle Lochkovian (Lower Devonian) conodonts and MAWSON, R. & TALENT, J. A., 1994, Age of an Early Devonian evolution of Flajsella n. gen. (Conodonta). 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America, Special Paper, 321: 131-144, 2 pls. (manuscriptreceivedDecember 9, 1998 W ANG, CHEN-YUAN & ZHANG, SHOU-AN, 1988, Discovery of che acceptedAugust26, 1999) early Early Devonian conodoncs from che Kuche area of Zinjiang: Journal of Scrarigraphy, 12 (2): 147-150, 4 pls [in Chinese] . Michael A. MuRPHY WILSON, G. A., 1989, Documentacion of conodont assemblages Deparrment of Geology, University of California a c Davis across che Lochkovian-Pragian (Early Devonian) boundary ac Davis, CA 95616, USA Wellington, Centrai New Souch Wales, Australia: Courier e-mail: [email protected] Forschungsinscicur Senckenberg, 117: 117-171, 12 pls. ZIEGLER, W., 1960, Conodonten aus dem Rheinischen Unrerdevon ]osé Ignacio VALENZUELA-Rros (Gedinnium) des Remscheider Sarrels (Rheinisches Deparramento de Geologia, Universicac de València Schiefergebirge): Palaonrologische Zeicschrift, 34: 169-20 l , 3 Dr. Moliner 50, E-46100 Burjassor, València, Spain pls. e-mail: [email protected]