Revision of Early Jurassic ammonoid types from the Per¸sani Mts ...

DOI:10.1127/0077-7749/2007/0243-0231 0077-7749/07/0243-0231 $ 6.00© 2007 E. Schweizerbart’sche Verlagsbuchhandlung, D-70176 Stuttgart

N. Jb. Geol. Paläont. Abh.2007, vol. 243/2, p. 231– 254, Stuttgart, February 2007, published online 2007

Revision of Early Jurassic ammonoid types from the Persani Mts.(East Carpathians, Romania)

Robert Tomas, Cluj and József Pálfy, Budapest

With 6 figures and 1 table

TOMAS, R. & PÁLFY, J. (2007): Revision of Early Jurassic ammonoid types from the Persani Mts.(East Carpathians, Romania). – N. Jb. Geol. Paläont. Abh., 243: 231– 254; Stuttgart.

Abstract: A large olistolith of Early Jurassic (Hettangian–Sinemurian) age exposed by the Tepei(Töpe) Creek near Racos (Alsórákos) (Persani Mts.) has been a prolific source of ammonoids.This locality furnished type specimens of 24 nominal species and 13 subspecies, introduced by sixdifferent authors between 1866 and 1953. A comprehensive revision of the type material is presented.The majority of the original specimens are available, two are believed lost and another five could notbe traced. Seven lectotypes are designated herein. Rediscovery of types for three species suppressestheir previously chosen neotypes. Included in this material are the type species of four ammonoidgenera. Confusion regarding the type species of Tragolytoceras is resolved by introducing T.bonarellii n. sp. and suggesting it as the new type species of the genus. Stratigraphic distribution ofsome of the revised taxa is assessed on the basis of new, detailed collections from a similar olistolithexposed a few km from the classical locality.

1. Introduction

The Tepei (Töpe) Creek near Racos (Alsórákos)(Romanian and Hungarian names, respectively) is oneof the classical Tethyan Early Jurassic ammonoidlocalities that has been known for 140 years. Therichly fossiliferous red nodular limestone and marlof “Rosso Ammonitico” or “Adnet” facies yielded awealth of specimens and furnished the type materialfor 37 nominal ammonoid taxa (species and sub-species), the revision of which forms the subject of thepresent study.

The locality was discovered by HERBICH (1866).His earliest collections were first studied by HAUER

(1866) who erected three new species (Ammonitesleptophyllus, A. transylvanicus, and A. altecinctus) onthe basis of HERBICH’s collection, without providingillustration. Later HERBICH (1878) published a mono-

graph that contained descriptions of 23 taxa, including10 new species. HERBICH’s work is illustrated by litho-graphic drawings that are in many cases idealized toan extent that hinders their use in identification of thetaxa. A new species of Lytoceras was proposed byBONARELLI (1900) partly on the basis of a specimenfigured by HERBICH (1878). VADÁSZ (1908) not onlyrevised HERBICH’s material but also significantlyaugmented the available paleontological informationbased on new collections from the area. In hismonograph 73 ammonoid taxa, including 8 newspecies and 9 subspecies, are described. TOULA

(1911) erected another species on the basis of hisstudy of private collections from the locality. LaterPREDA & RAILEANU (1953) studied the Liassic am-monoids from Mt. Persani and named one additionalnew species and 4 new subspecies. More recentlyPOPA & PATRULIUS (1996) summarized the occurrence

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232 R. Tomas and J. Pálfy

and biostratigraphy of ammonoid taxa from the area,including other localities discovered during thegeological mapping by the Geological Institute ofRomania.

Much of the collection of HERBICH and VADÁSZ,including many type specimens, was originally depo-sited in the Transylvanian National Museum (“ErdélyiNemzeti Múzeum”) in Kolozsvár or Cluj-Napoca(Hungarian and Romanian names, respectively). Thematerial was later transferred to the PaleontologicalMuseum of Babes-Bolyai University in Cluj-Napoca.RAKÚS (1993), doubting the existence of the typecollection, designated neotypes for two speciesoriginally described from Alsórákos. Four speciesin this assemblage were subsequently designatedas type species of newly introduced genera, docu-

mentation of their type material is especially im-portant.

We have studied the available collection at theBabes-Bolyai University of Cluj, where the majorityof the types are housed. Based on a comparison withpublished drawings and photographs, we attempted tolocate the type specimens for all taxa originallydescribed from Alsórákos. In only two cases oursearch has proved unsuccessful and we presume thatthose types are lost. The specimens studied by TOULA

(1911) were later, at least partly, incorporated into thecollections of the Geological Institute of Hungary,where his only type specimen has been identified.The originals of PREDA & RAILEANU (1953) werepresumably deposited in the Geological Institute ofRomania but, despite repeated enquiries, we have not

Fig. 1. Locality map. A– index map of the Persani Mts. inRomania; B – Index map of the region; C – Geographiclocation of the main outcrop.


been able to confirm their availability. The collectionsmay have been stored in a building that was seriouslydamaged in the 1977 Bucharest earthquake. It appearslikely that this material has been destroyed.

The aims of this study are (1) to provide a revisionand systematic treatment of the 37 nominal ammonoidspecies and subspecies, (2) report the availability orabsence (and likely loss) of the type specimens, (3)evaluate the status of types and formally designatelectotypes and paralectotypes where necessary, (4)give photographic illustration of the available typespecimens, (5) summarize the stratigraphic distri-bution from occurrences in a nearby section, and (6)discuss the status of four genera whose type speciesare based on this collection. A summary of the resultsof our revision is presented in Table 1.

2. Geological setting

The village of Racos (Alsórákos) lies along the OltRiver, near its gorge cutting through the Persani Mts.of the Eastern Carpathians. The Tepei (Romanianname, Töpe or Tepo in Hungarian) Creek is a south-north running tributary, entering the Olt 2.5 km east ofRacos (Fig. 1). The Persani Mts. are located SE of theTransylvanian basin in the central part of Romania,geographically attributed to the East Carpathians.Alsórákos is situated 60 km NW of Brasov (Brassó),the largest city in central Romania. The main fossili-ferous outcrops are situated 4 km SE from Alsórákosin the Töpe Valley.

The ammonite assemblage was collected from theLower Jurassic red nodular, “Ammonitico Rosso”-type limestone and marl. This lithofacies occurs asblocks of different size embedded as olistoliths in theBarremian-Aptian Wildflysch formation of the Buco-vinian Nappe, which forms part of the Transylvaniannappe system (PATRULIUS 1996).

Part of the collection originates from a large out-crop (referred to as the “main fossiliferous site” inHERBICH 1878 and VADÁSZ 1908) in the Töpe Valley.The other part was obviously collected from the creekbed as specimens weathered out and were transporteddownstream. These specimens are characteristicallyworn and often cannot be confidently identified.

Although the fauna is profusely documented inprevious studies, no stratigraphically controlled col-lection has been made before. Therefore the stratigra-phic distribution of the fauna has not been known.We recently carried out bed-by-bed collections froman outcrop of a similar olistolith exposed only a few

kilometers from the classical site. Thus it is nowpossible to establish the local stratigraphic range ofseveral taxa. Details of this study will be publishedseparately in the near future but reference is madehere to this new biostratigraphic information, unlessour recent collecting failed to yield additionalspecimens of a taxon.

3. Taxonomic revision

Taxa are listed and discussed here in the order of appear-ance in the original works, first referred to using theiroriginal bi- or trinomen. Specimen inventory numbers arethose currently used in the collection of PaleontologicalMuseum of Babes-Bolyai University. Synonymy lists arenot comprehensive and are generally confined to listingof topotypic material and, if available, a recent referencewhere a more comprehensive treatment of the taxon canbe found. Measurements are given in millimeters. Abbre-viations used are as follows: D = maximum diameter,U = umbilical diameter, WH = whorl height, WW = whorlwidth, WW/WH = ratio of whorl width and height,PRHW = number of primary ribs per half whorl.

3.1. Species erected by HAUER (1866)

The short account by HAUER (1866) contains only tersedescriptions and no illustration of three new species anda mention of one additional species identified fromHERBICH’s first collection. The specimens were returnedto HERBICH who later redescribed and figured the typematerial (HERBICH, 1878).

Ammonites leptophyllus HAUER, 1866Fig. 2a, b

Revised name: Geyeroceras leptophyllum (HAUER,1866).

Descriptions based on type material:1866 Ammonites leptophyllus n. sp. – HAUER, p. 192.1878 Phylloceras leptophyllum HAUER. – HERBICH, p.

112, pl. 20 H, Fig. 1a, b.1908 Phylloceras leptophyllum HAUER sp. – VADÁSZ, p.

342, Pl. 6, Fig. 7, 7a, Pl. 8, Fig. 1, 1a, Text-fig. 11.

Synonymy:1942 Geyeroceras leptophyllum HAUER. – KOVÁCS, p. 30.

(cum syn.)1953 Phylloceras leptophyllus HAUER. – RAILEANU (in

PREDA & RAILEANU), p. 342.1996 Geyeroceras leptophyllum (HAUER). – POPA &

PATRULIUS, pl. 16, figs. 8-9.

Type mater ia l : Holotype by monotypy is specimen No.12.571, a well-preserved phragmocone. The Cluj collectioncontains an additional 5 specimens, one of them is alsofigured in VADÁSZ (1908), but it appears from both HAUER

Revision of Early Jurassic ammonoid types from the Persani Mts. 233



Fig. 2 (Legend see p. 235)


(1866) and HERBICH (1878) that originally only onespecimen was available for description.

Measurements:

D U WH WW WW/WHHolotype 75.0 10.5 39.0 36.5 0.93

Remarks: KOVÁCS (1942) provided a detailed descriptionof this species and correctly assigned it to Geyeroceras. Asnoted in the original diagnosis, it is distinguished from G.cylindricum (SOWERBY) by its wider umbilicus and largersize. It also differs from G. subcylindricum (NEUMAYR)in a somewhat higher whorl expansion rate and in itstrapezoidal whorl cross section. The deep, conical umbilicusbears a vertical wall. It is also close to Calaiceras calais(MENEGHINI), but that form has well developed con-strictions on the phragmocone.

Ammonites transylvanicus HAUER, 1866Fig. 2g, h

Revised name: Juraphyllites transylvanicus (HAUER,1866).

Descriptions based on type material:1866 Ammonites transylvanicus n. sp. – HAUER, p. 192.1878 Phylloceras transylvanicum HAUER. – HERBICH, p.

114, pl. 20 J, fig. 1a, b. (non Pl. 20 H, Fig. 2a, b).1908 Rhacophyllites transylvanicus, HAU. sp. – VADÁSZ,

p. 323.

Synonymy:1901 Rhacophyllites transylvanicus HAUER. – FUCINI, p.

52, pl. 8, figs. 2-7.1996 Juraphyllites transilvanicus dorsoplanatus FUCINI.

– POPA & PATRULIUS, pl. 18, fig. 1.

Type mater ia l : Holotype by monotypy is specimenNo. 12.570T, a large body-chamber without phragmocone.

Measurements:


Remarks: The holotype is identified as the larger spe-cimen with missing phragmocone (substituted by a plastercast) figured by HERBICH (1878, pl. 20 I, fig. 1; note that the

drawing is idealized). Although HERBICH also illustratedan additional, smaller specimen (pl. 20 H, fig. 2, refiguredhere as Fig. 2c), HAUER’s original description refers to asingle specimen with the same diameter (4.5 inches, or c.11 cm). No suture lines can be observed, as only the bodychamber is preserved. The whorl shape and the bodychamber ornamentation bear juraphyllitid characters. Onlypart of the umbilicus is preserved; the umbilical wall, animportant feature in distinguishing juraphyllitid genera,cannot be observed on the missing phragmocone.

The body-chamber is characterized by flat flanks anddense ribbing. The ribs are straight on the external third ofthe flank where they are projected forward and cross theventer.

MEISTER & BÖHM (1993) suggested that this species issynonymous with J.nardii (MENEGHINI, 1853), which wouldhave priority. However, the body chamber of J. nardii ismore coarsely ribbed; the ribs are sigmoidal and arise nearthe umbilical wall. Although morphologically it appears adistinct species, curiously no other specimen has been foundin the collection or in our recent collections. HERBICH’sother figured specimen (No. 5890T) was refigured byVADÁSZ (1908, pl. 6, fig. 1) who correctly referred it toJ. gigas.

Ammonites altecinctus HAUER, 1866Fig. 2d, f

Revised name: Ectocetrites cf. petersi (HAUER, 1856).

Descriptions based on type material:1866 Ammonites altecinctus n. sp. – HAUER, p. 192.1878 Lytoceras altecinctum HAUER. – HERBICH, p. 116,

pl. 20 K, fig. 2.

Type mater ia l : Specimen No. 13.304, a large bodychamber fragment, is probably the holotype of the nominalspecies by monotypy and the original of HERBICH’sdrawing.

Measurements:

WH WW WW/WHHolotype 46.0 40.0 0.86

Remarks: HAUER (1866) described this species based on asingle body chamber fragment, later figured by HERBICH

(1878). The specimen has subquadratic whorl section withrounded margins. The ornamentation consists of closely


Fig. 2. Originals of HAUER (1866) and HERBICH (1878).a-b: Geyeroceras leptophyllum (HAUER, 1866), holotype, No. 12.571. a – lateral view, b – cross-section and ventralview, x1. c: Juraphyllites gigas (FUCINI, 1901), No. 5890T, original of HERBICH 1878, pl. 20H, fig. 2 (“Phyllocerastransylvanicum”), lateral view, x1. d, f: Ectocentrites cf. petersi (HAUER, 1856), holotype of “Ammonites altecinctusHAUER, 1866”, No. 13.304. d – lateral view, f – ventral view, x0.9. e: Angulaticeras marmoreum (OPPEL, 1862), holotypeof “Aegoceras tenuicostatum HERBICH, 1878”, No. 12.580, lateral view, x1. g-h: Juraphyllites transylvanicus (HAUER,1866), holotype, No. 12.570T. g – lateral view, h – ventral view, x0.9.



spaced and strong, radial ribs with small ventro-lateraltubercles. The ribs are projected forward at the ventro-late-ral shoulder. These characters suggest the body chamber ofan adult Ectocetrites petersi (HAUER, 1956), as alreadynoted by VADÁSZ (1908). WIEDMANN (1970) studied thespecimen but could not identify it with certainty withHERBICH’s original and labeled the specimen as Adnethi-ceras adnethicum (HAUER). However, A. adnethicum differsin its body chamber ornamentation of coarse annularribbing and well-defined ventral tubercles as well as in itselliptical whorl section.

The fragmentary nature of the type specimen renders“Ammonites altecinctus” a nomen dubium. Further con-fusion was created by SPATH (1924) who designatedLytoceras altecinctum the type species of his new genusTragolytoceras. The lack of phragmocone on the holotypeof type species makes it an unfortunate choice. Perhapsto circumvent this problem, or simply because of his un-familiarity with HERBICH’s specimen, SPATH clearly referredto BONARELLI’s specimen when proposing the new genus.The confusion of a genus based on a specimen taxono-mically different from the name-bearing type of the nominaltype species is resolved herein by proposing a new speciesfor BONARELLI’s specimen. For further discussion, seesection 4.4.

Strat ig raphic dis t r ibut ion: Late Hettangian, AngulataZone.

3.2. Species erected by HERBICH (1878)

Aegoceras althii HERBICH, 1878Fig. 3a, b

Revised name: Euagassiceras althii (HERBICH, 1878).

Descriptions based on type material:1878 Aegoceras Althii nov. sp. – HERBICH, p. 107, pl. 20.

E, fig. 1a, b.1908 Aegoceras Althii, HERB. – VADÁSZ, p. 371, pl. 9,

figs. 6-7.

Type mater ia l : Specimen No. 5917 appears the originalof HERBICH’s drawing and is considered as holotype byoriginal designation. It is said to be common but the numberof specimens available to HERBICH is not specified.

Measurements:


Remarks: This species, although common at this andother East Carpathian localities, has not yet been reportedfrom elsewhere. The type is a middle sized, evolutespecimen with subquadratic whorl cross section. The flanksare nearly flat. The venter is rounded, bearing a fine keelwithout sulci. The ornamentation consists of strong, radialribs on the flanks, bearing a small tubercle on the ventro-lateral shoulder where they are projected forward and reachthe keel. This form appears closely related to E. terquemi(REYNES) (see e.g. CORNA et al. 1997). It also resemblesE. sauzeanum but differs in being more evolute and in itslower whorl expansion rate.

Strat ig raphic dis t r ibut ion: Early Sinemurian, Semi-costatum Zone.

Aegoceras alutae HERBICH, 1878Fig. 3e, f, i, j

Revised name: Ectocentrites? alutae (HERBICH, 1878).

Descriptions based on type material:1878 Aegoceras Alutae nov. sp. – HERBICH, p. 109, pl.

20 B, fig. 3a, b.1908 Ectocentrites Petersi, HAU. sp. – VADÁSZ, p. 357,

pl. 10, fig. 1.

Type mater ia l : HERBICH (1878) reported three spe-cimens, of which No. 13.340 is the figured original in thefirst description (and also in VADÁSZ (1908)), thereforeconsidered as the holotype by original designation.

Measurements:

D U WHHolotype 60 30 17

Remarks: The type specimen is complete but poorlypreserved. The shell is evolute with elliptical whorl section.The ornamentation is characterized by somewhat irregularlyspaced, gently swaying, wiry ribs with well-developedtubercles (or small spines on the other better preserved

Fig. 3. Originals of HERBICH (1878).a-b: Euagassiceras althii (HERBICH, 1878), holotype, No. 5917. a – lateral view, b – ventral view, x1. c-d: Arietitidae indet.,lectotype of “Aegoceras Albense HERBICH, 1878”, No. 5910.H. c – lateral view, d–ventral view, x1. e-f, i-j: Ectocentrites?alutae (HERBICH, 1878), holotype, No. 13.340. Photographed specimen, e – lateral view, f – ventral view, x1. Drawingsreproduced from HERBICH’s illustration, i – lateral view, j – ventral view, x1. Note difference of drawing from actualspecimen; such idealized rendering is typical of illustrations in HERBICH (1878). g-h: Geyeroceras cylindricum (SOWERBY,1831), holotype of “Phylloceras bielzii HERBICH, 1878”, No. 5798.H. g – lateral view, h – cross-section and ventral view, x1.k-l: Phylloceras persanense (HERBICH, 1878). k – paralectotype, No. 5893.H, lateral view, x1; l – lectotype, No. 12.575.H,lateral view, x1.



specimens) on the ventral shoulder. Inner whorls bear moredensely spaced and lower ribs, and sparse constrictions.

WÄHNER (1882) noted that HERBICH’s form couldn’t bedistinguished from Ectocentrites petersi, a view later sharedby VADÁSZ (1908). The juvenile stage of both species ischaracterized by constrictions and densely ribbed innerwhorls with ventral tubercles. However, the sub-adultspecimens (50–60 mm in diameter) are distinguished bytheir whorl cross-section and ornamentation. The ellipticalwhorls of E.? alutae differ from sub-rectangular whorlsection of E. petersi, which is also more densely ribbed withribs that cross over the venter and bear tubercles that aremuch smaller than those of E. alutae. Assignment to Ecto-centrites is probable, although the tuberculate ventral ribtermination may be interpreted a distinctive character evenat the genus level.

The type specimen as figured in HERBICH (1878) isa typical example for the idealization common in hisdrawings. For comparison, we reproduce the originalillustration (Fig. 3i, j) along the photograph of the specimen(Fig. 3e, f).


Aegoceras tenuicostatum HERBICH, 1878Fig. 2e

Revised name: Angulaticeras marmoreum (OPPEL,1862).

Descriptions based on type material:1878 Aegoceras tenuicostatum nov. sp. – HERBICH, p.

110, pl. 20D, fig. 3a, b.1908 Schlotheimia marmorea, OPP. sp. – VADÁSZ, p. 363.

Synonymy:1988 Angulaticeras marmoreum (OPPEL). – BLOOS, p. 6,

pls. 1-3, pl. 4, figs. 7-8, pl. 5-8, pl. 9, fig. 1 (cumsyn.)

1993 Angulaticeras marmoreum (OPPEL). – RAKÚS &LOBITZER, p. 926, pl. 2, fig. 2.

1999 Angulaticeras marmoreum (OPPEL, 1862). – BÖHM

et al., p. 188, pl. 26, fig. 12000 Angulaticeras marmoreum (OPPEL). – SMITH &

TIPPER, p. 4, Pl. 1, Figs. 1-3.

Type material: Specimen No. 12.580 appears theoriginal of HERBICH’s drawing and considered as holo-type of the nominal species by original designation.

Measurements:

D U WH WW WW/WHHolotype 43.8 11 19 12.8 0.67

Remarks: The type and other specimens in the collectionare confidently assigned to Angulaticeras marmoreum, asalready noted by VADÁSZ (1908) and confirmed by BLOOS

(1988), who provides a detailed revision of this widelydistributed species.


Aegoceras albense HERBICH, 1878Fig. 3c, d

Revised name: Arietitidae indet.

Descriptions based on type material:1878 Aegoceras Albense nov. sp. – HERBICH, p. 111, pl.

20A, fig. 3a, b.1908 Aegoceras albense HERB. – VADÁSZ, p. 371.

Type mater ia l : Specimen No. 5910.H, with 26 mm in itslargest dimension, is the original of HERBICH’s drawing anddesignated here as lectotype of the nominal species.

Remarks: The figured specimen is an indeterminableinner whorl fragment. VADÁSZ (1908) commented on itsunsuitability as a type and did not illustrate it, neverthelessaccepted the species. The preserved sutures suggest anarietitid but a more precise identification is not possible. Werecommend abandoning this name.

Phylloceras persanense HERBICH, 1878Fig. 3k, l

Revised name: Phylloceras persanense HERBICH, 1878.

Descriptions based on type material:1878 Phylloceras Persanense nov. sp. – HERBICH, p.

111, pl. 20E, fig. 3a, b, pl. 20F, fig. 1.1908 Phylloceras persanense HERB. – VADÁSZ, p. 341,

pl. 7, fig. 1, 1a, text-fig. 10.

Synonymy:1900 Phylloceras persanense HERBICH. – UHLIG, p. 15,

pl. 1, fig. 1a, b, text-fig. 1.1901 Phylloceras persanense HERBICH. – FUCINI, p. 22,

pl. 4, figs. 1-2.1908 Phylloceras infraliasicum, nov. sp. – VADÁSZ, p.

316, pl. 9, fig. 2, 2a, text-fig. 17.1953 Phylloceras persanense HERB. – RAILEANU (in

PREDA & RAILEANU, 1953), p. 341, pl. 4, fig. 2, 2a.1996 Geyeroceras persanense (HERBICH). – POPA &

PATRULIUS, pl. 17, figs. 3, 6.

Type mater ia l : HERBICH’s first description was based onseveral specimens, of which two were illustrated. The largerone, No. 12.575.H, is designated herein lectotype, whereasthe smaller one, No. 5893.H is a paralectotype. Othermembers of the original type series cannot be identifiedwith certainty, although altogether thirteen specimens existnow in the collection. The types are well-preserved internalmoulds of phragmocones.


Measurements:

D U WH WW WW/WHLectotype 111.0 12.3 61.8 41.8 0.67Para-lectotype 97.5 10.5 54.6 40.0 0.73

Remarks: On the flanks of the better-preserved specimens(including the lectotype) vague, irregular constrictionsappear that are strongly exaggerated and falsely shortenedon the drawing of HERBICH (1878). The maximum whorlwidth is attained at mid-flank rather than at the umbilicalshoulder as shown by HERBICH.

WIEDENMAYER (1977) regarded this species a repre-sentative of Calaiceras, a genus that mainly includes Pliens-bachian forms characterized by indistinct constrictions,variable in size and number (GÉCZY & MEISTER 1998). Itcannot be convincingly demonstrated that distinctivegeneric characters are beyond the range of intraspecificvariability. Therefore the conservative approach of assign-ment to Phylloceras is preferred here.

A specimen described by VADÁSZ (1908) as a newspecies, P. infraliasicum is regarded a junior synonym ofP. persanense. Its extremely narrow umbilicus, the allegeddiagnostic feature, is a preservational artifact of the stronglydeformed phragmocone.

Strat ig raphic dis t r ibut ion: Late Hettangian.

Phylloceras bielzii HERBICH, 1878Fig. 3g, h

Revised name: Geyeroceras cylindricum (SOWERBY,1831).

Descriptions based on type material:1878 Phylloceras Bielzii nov sp. – HERBICH, p. 113, Pl.

20 F, fig. 2a, b.1908 Phylloceras cylindricum, SOW., var. Bielzii, HERB. –

VADÁSZ, p. 340, text-fig. 9.

Synonymy: 1878 Phylloceras cylindricum SOWERBY. – HERBICH, p.

113, pl. 20E, fig. 2a, b.1908 Phylloceras cylindricum, SOW. sp. – VADÁSZ; p.

338, text-fig. 7.1953 Phylloceras cylindricum SOW. var. bielzii HERB. –

RAILEANU (in PREDA & RAILEANU, 1953), p. 342,pl. 2, fig. 2a, b, c.

1993 Geyeroceras cylindricum (SOWERBY, 1831). –RAKÚS & LOBITZER, p. 923, pl. 2, fig 1. (cum syn.).

1996 Geyeroceras cylindricum bielzii HERBICH. – POPA

& PATRULIUS, pl. 17, fig. 1.1999 Geyeroceras cylindricum (SOWERBY, 1831). –

BÖHM et al., p. 186, pl. 25, fig. 6. (cum syn.).

Type material: Specimen No. 5798.H is holotype ofthe nominal species by original designation, theoriginal of HERBICH’s drawing, a poorly preservedinternal mould of an incomplete body-chamber.

Measurements:


Remarks: The flat flanks, quadratic whorl section, andnarrow umbilicus of this form agree well with Geyerocerascylindricum. Already VADÁSZ (1908) included it in thisspecies as a variety. HERBICH regarded the narrow umbilicusa distinguishing feature, but it is exaggerated on the drawingand it falls within the variability of G. cylindricum. Otherslight differences, the somewhat higher whorl section andlarger size do not warrant maintaining it as a separate spe-cies or subspecies.

Strat ig raphic dis t r ibut ion: Hettangian to Sinemurian.

Phylloceras ürmösense HERBICH, 1878Fig. 4a

Revised name: Paradasyceras uermoesense (HERBICH,1878).

Descriptions based on type material:1878 Phylloceras Ürmösense nov. sp. – HERBICH, p. 113,

pl. 20K, fig. 1a, b.1908 Rhacophyllites ürmösensis, HERB. sp. – VADÁSZ, p.

330, pl. 6, fig. 5, text-fig. 4.

Synonymy: 1953 Rhacophyllites (Paradasyceras) ürmösensis HERB.

– RAILEANU (in PREDA & RAILEANU), p. 340, pl. 1,fig. 1, pl. 2, fig. 1.

1953 Rhacophyllites (Paradasyceras) ürmösensis HERB.var. tenuilobata (nov. var.). – RAILEANU (in PREDA

& RAILEANU), p.340, pl.3, fig.1; pl.5, fig. 2.1993 Paradasyceras uermoesense (HERBICH, 1878). –

RAKÚS, p. 948, Pl. 4, figs. 1-2, 4. (cum syn.)1996 Paradasyceras uermoesense HERBICH. – POPA &

PATRULIUS, pl. 16, fig. 1, pl. 17, fig. 2, pl. 18, Fig. 3.1996 Paradasyceras tenuilobata RAILEANU. – POPA &

PATRULIUS, pl. 16, fig. 2.

Type material: Specimen 5897.H is the original ofHERBICH’s and VADÁSZ’s figure, is considered asholotype by original designation.Another 7 specimensexist in the collection, some may have been part of theoriginal type series.

Measurements:


Remarks: The holotype is a phragmocone, but otherspecimens from the collection display smooth body cham-ber with no growth lines. The diagnostic features aresummarized by RAKÚS (1993) who designated a neotype ashe doubted the existence of HERBICH’s original. According



to the ICZN (1999, Art. 75.8), our rediscovery of theholotype invalidates the chosen neotype. SPATH (1923)designated this species as type species of Paradasyceras,see further discussion in section 4.3. Umbilical width varieswithin the species, therefore distinguishing the subspeciesP. uermoesense tenuilobata RAILEANU is not supported(see below: Rhacophyllites (Paradasyceras) ürmösensisHERBICH tenuilobata RAILEANU, 1953).


Phylloceras rákosense HERBICH, 1878Fig. 4d

Revised name: Juraphyllites rakosensis (HERBICH,1978).

Descriptions based on type material:1878 Phylloceras Rákosense nov. sp. – HERBICH, p. 114,

pl. 20G, fig. 3a, b, c.1908 Rhacophyllites rákosensis HERB. sp. – VADÁSZ, p.

328, pl. 6, fig. 3.

Type material: The holotype, original of HERBICH’sfigure, is specimen No. 5811.H, a complete but some-what worn internal mould.

Measurements:


Remarks: The specimen is medium-sized, laterally com-pressed, mid-volute with rapidly expanding whorls andvertical umbilical wall. Its distinctive feature is the bodychamber ornamentation: widely spaced, sigmoidal ribs thatarise at the umbilical edge and are the strongest on lowerthird of flank. The venter bears fine, intercalated secondaryribs, their faintness may be an artifact of abrasion. Notethat HERBICH’s drawing does not render the ornamentationfaithfully. Sutural pattern, volution and ornamentationshow juraphyllitid characters. The taxon is type species ofDasyceras HYATT, 1900, and the status of this genus isdiscussed in section 4.1.

Phylloceras aulonotum HERBICH, 1878Fig. 4b

Revised name: Schistophylloceras aulonotum (HERBICH,1878).

Descriptions based on type material:1878 Phylloceras aulonotum nov. sp. – HERBICH, p. 115,

pl. 20G, fig. 2a, b, c.1908 Rhacophyllites (Kochites) aulonotus, HERB. sp. –

VADÁSZ, p. 336, pl. 6, fig. 6, text-fig. 6.

Synonymy:1953 Rhacophyllites (Schistophylloceras) aulonotum

HERB. – RAILEANU (in PREDA & RAILEANU), p. 341,pl. 4, fig. 1.

1993 Schistophylloceras aulonotum (HERBICH, 1878). –RAKÚS, p. 949, pl. 3, fig. 1. (cum syn.).

Type material: The species is said to be common byHERBICH (1878). The holotype is specimen No.2055.H, the original of figures by both HERBICH andVADÁSZ, a well-preserved internal mould with part ofthe body chamber. One additional specimen exists inthe collection.

Measurements:


Remarks: The main diagnostic feature is the deep, pro-minent ventral furrow on the internal mould. Other charac-teristics of the species are summarized by RAKÚS (1993)who designated a neotype as he doubted the existence ofHERBICH’s original. According to the ICZN (1999, Art.75.8), our rediscovery of the holotype invalidates the chosenneotype. The taxon is type species of SchistophyllocerasHYATT, 1900, and the status of this genus is discussed insection 4.2.

Strat ig raphic dis t r ibut ion: Early to Late Hettangian.


Fig. 4. Originals of HERBICH (1878), BONARELLI (1900) and VADÁSZ (1908).a: Paradasyceras uermoesense (HERBICH, 1878), holotype, No. 5897.H, lateral view, x1. b: Schistophylloceras aulonotum(HERBICH, 1878), holotype, No. 2055.H, lateral view, x1. c: Tragolytoceras bonarellii nov. sp., holotype, Natural HistoryMuseum (London) No. C.8480, from Pontalto (Italy), lateral view, x1. d: Juraphyllites rakosensis (HERBICH, 1878),holotype, No. 5811.H, lateral view, x1. e-f: Calliphylloceras nilssoni (HEBERT, 1866), holotype of “Phylloceras sylvestreHERBICH, 1878”, No. 12.573.H. e – lateral view, f – cross-section and ventral view, x1. g-h: Juraphyllites gigas (FUCINI,1901), holotype of “Rhacophyllites gigas intermedia VADÁSZ, 1908”, No. 5810.H. g – lateral view, h – cross-sectionand ventral view, x1. i -j: Geyeroceras cylindricum (SOWERBY, 1831), holotype of “Phylloceras szádeczkyi VADÁSZ, 1908”,No. 2052.H. i – lateral view, j – cross-section and ventral view, x1. k-l: Phylloceras psilomorphum (NEUMAYR, 1879),types of “Phylloceras hungaricum VADÁSZ, 1908”. k – lectotype, No. 5886(a), lateral view, x1; l – paralectotype,No. 5886(b), lateral view, x1.


Phylloceras sylvestre HERBICH, 1878Fig. 4e, f

Revised name: Calliphylloceras nilssoni (HEBERT,1866).

Descriptions based on type material:1878 Phylloceras sylvestre nov. sp. – HERBICH, p. 115, pl.

20G, fig. 1a, b, c.1908 Phylloceras sylvestre, HERB. – VADÁSZ, p. 354, pl.

7, fig. 5, 5a, text-fig. 22.

Synonymy:1996 Calliphylloceras sp., aff. C. sylvestre (HERBICH). –

POPA & PATRULIUS, pl. 16, fig. 6.2000 Calliphylloceras nilssoni (HEBERT, 1866). – JOLY,

p. 85, pl. 19, figs. 6-9, text-figs. 177-181 (cumsyn.).

Type material: Holotype of the nominal species bymonotypy is specimen 12.573.H.

Measurements:


Remarks: An internal mould of phragmocone withnarrow umbilicus and constrictions, both features aremisrepresented on the drawing of HERBICH. VADÁSZ

(1908), although accepted the species, emphasized itssimilarity to Phylloceras nilssoni. JOLY (2000) recentlydiscussed this species with no mention to the Transyl-vanian material. Nevertheless his description leaves littledoubt that HERBICH’s specimen can be assigned to thistaxon.

Strat ig raphic dis t r ibut ion: Toarcian, Bifrons Zone.

3.3. Species erected by BONARELLI (1900)

Lytoceras herbichi BONARELLI, 1900Fig. 5b, c

Revised name: Adnethiceras herbichi (BONARELLI,1900).

Description based on the type material:1878 Aegoceras adnethicum HAUER. – HERBICH, p. 108,

pl. 20 C, fig. 1a, b, c.1908 Aegoceras adnethicum HAU. sp. var. involuta, nov.

var. – VADÁSZ, p. 368, pl. 9, fig. 4, 4a, text-fig. 24.1900 Lytoceras Herbichi n. f. – BONARELLI, p. 69.

Synonymy: 1908 Aegoceras simplex nov. sp. – VADÁSZ, p. 370, pl. 9,

fig. 5, 5a.1953 Lytoceras (Tragolytoceras) herbichi BONAR. –

RAILEANU (in PREDA & RAILEANU), p. 345, pl. V,fig. 1, 1a.

1970 Adnethiceras herbichi (BONARELLI). – WIEDMANN,p. 1000, pl. 9, fig. 1, text-fig. 30P (cum syn.).

Type material: Holotype by monotypy is specimenNo. 12.578.H, as stated by WIEDMANN (1970). Theoriginal of HERBICH (1878) is a well-preserved phrag-mocone with a partly preserved body-chamber.

Measurements:


Remarks: HERBICH (1878) described and figured a singlespecimen as Aegoceras adnethicum. BONARELLI (1900)proposed it as a new species, distinguished from A.adnethicum by its denser ribbing and higher whorl ex-pansion rate. WIEDMANN (1970) provides a review of thegenus Adnethiceras and retains A. herbichi as a separatespecies. See also discussion below, on taxa Aegocerasadnethicum HAUER var. involuta VADÁSZ, 1908 and Aego-ceras simplex VADÁSZ, 1908.

3.4. Species and subspecies erected byVADÁSZ (1908)

Rhacophyllites gigas FUCINI intermedia VADÁSZ,1908Fig. 4g, h

Revised name: Juraphyllites gigas (FUCINI, 1901).

Description based on type material:1908 Rhacophyllites gigas FUC. var. intermedia nov. var.

– VADÁSZ, p. 327, pl. 6, fig. 2, 2a, text-fig. 3.


Fig. 5. Originals of VADÁSZ (1908).a: Zetoceras prinzi (VADÁSZ, 1908), holotype, No. 5884.H, lateral view, x1. b-c: Adnethiceras herbichi (BONARELLI, 1900),holotype, No. 12.578.H, (also the lectotype of Aegoceras adnethicum involuta VADÁSZ, 1908”), HERBICH’s collection.b – lateral view, c – ventral view, x1. d-e: Zetoceras prinzi (VADÁSZ, 1908), lectotype of “Phylloceras oenotrium com-planata VADÁSZ, 1908”, No. 2047.H. d – cross-section and ventral view, e – lateral view, x1. f: Adnethiceras herbichi(BONARELLI, 1900), holotype of “Aegoceras simplex VADÁSZ, 1908”, No. 5911.H, lateral view, x1. g-h, j-k: Echioceras?raricostatoides (VADÁSZ, 1908), paralectotype, No. 5918(b), g – lateral view, h – ventral view, x1; lectotype, No. 5918(a),j – ventral view, k – lateral view, x1. i: Phylloceras persanense (HERBICH, 1878), holotype of “Phylloceras infraliasicumVADÁSZ, 1908”, No. 5885.H, lateral view, x1.



Synonymy:1901 Rhacophyllites gigas n. sp. – FUCINI, p. 56, pl. 9,

figs. 2-5, text-fig. 27.1953 Rhacophyllites gigas FUC. – RAILEANU (in PREDA

& RAILEANU), p. 338, pl. 3, fig. 2.1953 Rhacophyllites gigas FUC. var. intermedia VAD. –

RAILEANU (in PREDA & RAILEANU), p. 338, pl. 1,fig. 4.

Type material: Specimen No. 5810.H, a phragmocone,is holotype of the nominal subspecies by monotypy.

Measurements:


Remarks: The shell is convolute, laterally compressed, thewhorl cross-section is elliptical and the flanks are convexwith rounded venter. The suture line is typically jura-phyllitid. The body-chamber is not preserved, hence itsornamentation cannot be observed. The morphologicalfeatures of our specimen are in good agreement withthe original description of Juraphyllites gigas (FUCINI,1901; pl. 11, figs. 2-5), although with a somewhat smallerumbilicus.

Phylloceras szádeczkyi VADÁSZ, 1908Fig. 4i, j

Revised name: Geyeroceras cylindricum (SOWERBY,1831).

Descriptions based on type material:1908 Phylloceras Szádeczkyi nov. sp. – VADÁSZ, p. 344,

pl. 7, fig. 2, 2a, text-fig. 12.

Synonymy:1996 Geyeroceras szadeczkyi (VADÁSZ). – POPA &

PATRULIUS, pl. 16, fig. 5.1999 Geyeroceras cylindricum (SOWERBY, 1831). –

BÖHM et al., p. 186, pl. 25, fig. 6. (cum syn.).

Type material: Specimen No. 2052.H, the original ofVADÁSZ (1908), is the holotype by monotypy.

Measurements:


Remarks: VADÁSZ (1908) notes similarity to other speciesnow accommodated in the genus Geyeroceras. There areonly minor differences compared to the original diagnosisof the type species of G. cylindricum. The flanks are nearlyflat with a small convexity, the venter is rounded; the whorlcross-section is higher, the umbilicus wider. VADÁSZ’s typebest agrees with the original of FUCINI’s (1901) subspeciesG. cylindricum compressa.


Phylloceras lipoldi HAUER primitiva VADÁSZ, 1908

Original description:1908 Phylloceras Lipoldi HAU. sp. var. primitiva nov. var.

– VADÁSZ, p. 347, pl. 8, fig. 2, 2a, text-fig. 14.

Type material: The type specimen of this subspecieswas not found in the collection.

Phylloceras hungaricum VADÁSZ, 1908Fig. 4k, l

Revised name: Phylloceras psilomorphum (NEUMAYR,1879).

Descriptions based on type material:1908 Phylloceras hungaricum nov. sp. – VADÁSZ, p. 348,

pl. 8, figs. 3, 3a, 4, 4a, text-fig. 16.

Synonymy:1879 Phylloceras psilomorphum n. f. – NEUMAYR, p. 21,

pl. 2, fig. 4.1999 Phylloceras psilomorphum (NEUMAYR, 1879). –

BÖHM et al., p. 185, pl. 24, fig. 4, text-fig. 36. (cumsyn.).

Type material: Both specimens of the original typeseries exist, under a single inventory number (No.5886.H). The larger one is designated here as lecto-type, and the smaller one as paralectotype.

Measurements:

D U WH WW WW/WHLecto-type 52.0 7.2 28.6 19.6 0.68Para-lectotype 42.5 5.3 24.8 13.0 0.52

Remarks: VADÁSZ‘s types are in good agreement with theoriginal description of P. psilomorphum. The cross sectionof the whorls is somewhat wider and the umbilicus isnarrower in comparison with the lectotype figured byNEUMAYR (1879). We consider the two forms synonymous,and the small differences fall within the range of morpho-logical variation of this species.

Strat ig raphic dis t r ibut ion: Late Hettangian.

Phylloceras infraliasicum VADÁSZ, 1908Fig. 5i

Revised name: Phylloceras persanense (HERBICH,1878).


Descriptions based on type material:1908 Phylloceras infraliasicum nov. sp. – VADÁSZ; p.

350, Pl. IX, Fig. 2, 2a, Text-fig. 17.

Synonymy: see above under taxon Phylloceras persa-nense HERBICH, 1878.

Type material: Specimen No. 5885.H is holotype ofthe nominal species by monotypy, a poorly preserved,strongly deformed and laterally compressed internalmould of phragmocone.

Measurements:

D U WH WW WW/WHHolotype 98,5 4,0 64,2 ~ 25,5 0,39

Remarks: P. infraliasicum is regarded as a junior synonymof P. persanense (see discussion above), their apparentminor difference is explained by deformation of the typespecimen of the former species.

Phylloceras prinzi VADÁSZ, 1908Fig. 5a

Revised name: Zetoceras prinzi (VADÁSZ, 1908).

Descriptions based on type material:1908 Phylloceras Prinzi, nov. var. – VADÁSZ, p. 351, pl.

8, fig. 5, 5a, text-fig. 18.

Synonymy:1908 Phylloceras oenotrium FUC. sp. var. complanata,

nov. var. – VADÁSZ, p. 352, pl. 8, fig. 6, 6a, text-fig. 20.

1999 Zetoceras complanatum (VADÁSZ, 1908). – BÖHM

et al., p. 186, pl. 24, fig. 2.

Type material: Holotype by monotypy is the specimenNo. 5884.H, a poorly preserved phragmocone.

Measurements:


Remarks: The shell is involute, laterally compressed. Thewhorls have flat flanks and triangular whorl cross-section,rounded venter and umbilical margin. The whorl cross-section attains the largest width at the umbilical margin.Another similar form described by VADÁSZ as a new sub-species Phylloceras oenotrium complanata is consideredsynonymous with this species. See further discussion underthis taxon.

Phylloceras oenotrium FUCINI var. complanataVADÁSZ, 1908

Fig. 5d, e

Revised name: Zetoceras prinzi (VADÁSZ, 1908).

Descriptions based on type material:1908 Phylloceras oenotrium FUC. var. complanata nov.

var. – VADÁSZ, p. 352, pl. 8, fig. 6, 6a, text-fig.20.

Synonymy:1908 Phylloceras Prinzi nov. sp. – VADÁSZ, p. 351, pl. 8,

fig. 5, 5a, text-fig. 18.1953 Phylloceras oenotrium FUCINI var. complanata

VAD. – RAILEANU (in PREDA & RAILEANU), p. 343,pl. 3, fig. 3, 3a.

1999 Zetoceras complanatum (VADÁSZ, 1908). – BÖHM

et al., p. 186, pl. 24, fig. 2. (cum syn.).

Type material: There are two internal moulds withpreserved phragmocone under the same inventorynumber (2047.H) in VADÁSZ’s collection. The larger,originally figured specimen is designated here aslectotype of the nominal subspecies.

Measurements:

D U WH WW WW/WHLecto-type 51.8 7.2 27.6 19.8 0.66

Remarks: The subspecies Phylloceras oenotrium com-planata was accepted and elevated to species rank byRAKÚS (in BÖHM et al., 1999) on the basis of differences inthe whorl cross-section from Zetoceras oenotrium (FUCINI).We consider this taxon as a junior synonym of Z. prinzi(VADÁSZ) that takes priority by virtue of being describedfirst in the same reference (on. p. 317 in VADÁSZ, 1908,whereas Z. oenotrium complanata appears on p. 318). Seealso discussion of taxon Phylloceras prinzi VADÁSZ, 1908.

Aegoceras adnethicum HAUER var. involutaVADÁSZ, 1908

Fig. 5b, c


Description based on the type material:1878 Aegoceras adnethicum HAUER – HERBICH, p. 108,

pl. 20C, fig. 1a, b, c.1908 Aegoceras adnethicum HAU. sp. var. involuta nov.

var. – VADÁSZ, p. 368, pl. 9, fig. 4, 4a, text-fig. 24.

Synonymy:1886 Aegoceras adnethicum v. HAU. – GEYER, p. 261, pl.

4, figs. 2-3.1900 Lytoceras Herbichi n. f. – BONARELLI, p. 69.1970 Adnethiceras Herbichi (BONARELLI) – WIEDMANN,

p. 1000, pl. 9, fig. 1, text-fig. 30P (cum syn.).

Type material: Specimen No. 12.578.H, the original ofboth HERBICH’s drawing and VADÁSZ’s photograph, is



a well preserved phragmocone with partly preservedbody-chamber. It is designated here as lectotype forthis nominal subspecies, of which VADÁSZ had onefragmentary and two complete specimens.

Measurements: See under taxon Lytoceras herbichiBONARELLI, 1900.

Remarks: VADÁSZ (1908) proposed the new subspecieson the basis of the following distinctive features: asmaller umbilicus (U/D = 0.42 vs. 0.51), more denseribbing (PRHW = 40 vs. 33), slightly compressed vs.slightly depressed semicurcular whorl cross-section, and alack of nodes on the ring-like ribs. WIEDMANN (1970)de-emphasizes the differences and retains A. adnethicumand A. herbichi as closely related species. We regardAegoceras adnethicum HAUER var. involuta VADÁSZ, 1908as a junior synonym of Adnethiceras herbichi (BONARELLI,1900). See also discussion of taxon Lytoceras herbichiBONARELLI, 1900.

Aegoceras simplex VADÁSZ, 1908Fig. 5f


Description based on the type material:1908 Aegoceras simplex nov. sp. – VADÁSZ, p. 370, pl. 9,

fig. 5, 5a, text-fig. 25.

Synonymy: Same as for taxon Aegoceras adnethicumHAUER var. involuta VADÁSZ, 1908.

Type material: Specimen No. 5911.H is holotype bymonotypy of the nominal species, a poorly preservedphragmocone.

Measurements:


Remarks: Aegoceras simplex (VADÁSZ, 1908) is regardedhere as a junior synonym of Adnethiceras herbichi

(BONARELLI, 1900), although WIEDMANN (1970) tentativelyretained it as separate species of Adnethiceras. See alsodiscussion of taxon Lytoceras herbichi BONARELLI,1900.

Arietites raricostatoides VADÁSZ, 1908Fig. 5g, h, j, k

Revised name: Echioceras? raricostatoides (VADÁSZ,1908).

Descriptions based on type material:1908 Arietites raricostatoides nov. sp. – VADÁSZ, p. 373,

text-fig. 26.

Synonymy:1973 Echioceras raricostatoides (VADÁSZ). – GETTY, pl.

1, fig. 12.1993 Echioceras raricostatoides VADÁSZ. – DOMMER-

GUES, p. 134, pl. 7, fig.2.1998 Echioceras raricostatoides VADÁSZ, 1908. – BLAU,

p. 206, pl. 4, figs. 3-8 (cum syn.).

Type material: Two specimens under inventory No.5918 are assumed the originals of VADÁSZ (1908). Thelarger one is designated here as lectotype, the smallerone paralectotype. No illustration is given by VADÁSZ

(1908), thus identification of the types is necessarilybased on the description only.

Measurements:

D U WH WW WW/WHLectotype 73.8 43.0 20.0 16.5 0.82Para-lectotype 61.0 37.5 13.8 10.0 0.72

Remarks: The available specimens are rather poorlypreserved internal moulds, observing all morphologicalfeatures remains difficult. The lectotype has a long bodychamber of at least 7/8 whorl, bears a faint keel and sparseribs that are confined to the flanks and fade out on venter.A fragment of an inner whorl reveals more dense ribbingin early ontogeny. The paralectotype is cut in half to showsubcircular whorl section, its outer three whorls are exposedbut the inner ones are not visible up to D = 1.3 cm. It bears


Fig. 6. Originals of VADÁSZ (1908) and TOULA (1911).a-b, d, h: Euagassiceras spinaries (QUENSTEDT, 1858), syntypes of “Arietites obtusus vulgaris VADÁSZ, 1908”, No.12.579H(a), a – lateral view, b – ventral view, x1; No. 5916.H, d – lateral view, x1; No. 12.579H(b), h – lateral view, x1.c: Coroniceras? sp., holotype of “Arietites pseudospiralis VADÁSZ, 1908”, No. 5920.H, lateral view, x0.9. e: Coronicerascf. rotiforme (SOWERBY, 1824), holotype of “Arietites subrejectus VADÁSZ, 1908”, No. 5921.H, lateral view, x1. f: Coroni-ceras cf. rotiforme (SOWERBY, 1824), holotype of “Arietites carenatus antiqua VADÁSZ, 1908”, No. 5919.H, lateral view, x1.g: Caenisites plotti (REYNES, 1879), holotype of “Arietites altesulcatus involuta VADÁSZ, 1908”, No. 5924.H, lateralview, x1. i: Lytoceras ‘lexeni’ (TOULA, 1911), lectotype, Hungarian Geological Institute No. J.11475, lateral view, x1.j: Metophioceras? simplex (VADÁSZ, 1908), holotype of “Arietites spiratissimus simplex VADÁSZ, 1908”, No. 5927.H,lateral view, x1.



a weak keel, and is sparsely but regularly costate. BUCKMAN

(1914) was the first to question the existence of the originalsof VADÁSZ (1908) and designated a lectotype from theBayle collection obtained at Seichamp near Nancy, France(GETTY 1973), used in subsequent identification of thespecies. GETTY (1973) was unable to find BUCKMAN‘slectotype and he selected a neotype from the samecollection. That specimen was accepted as a neotype byDOMMERGUES (1993) but later BLAU (1998) still regardsVADÁSZ’s specimen as holotype. Most authors distinguishthe species raricostatoides from raricostatum, the firstrepresenting morphologies with sparse ribbing and roundedwhorl cross-section. This species is of stratigraphical im-portance as the name-bearing taxon of the Raricostatoides(Sub)zone within the Late Sinemurian (see discussion inDEAN et al. 1961). The rediscovery of the lectotype invali-dates previous neotype selections and will allow for areappraisal of the frequently reported northwest Europeanform identified with E. raricostatoides. The generic attri-bution is questionable as the preservation is mediocre andthe Upper Sinemurian is not well represented in the typearea, therefore homeomorphic other Early Jurassic genera(e.g. Caloceras, Epophioceras) cannot be excluded.

Arietites obtusus SOWERBY var. vulgaris VADÁSZ

Fig. 6a, b, d, h

Revised name: Euagassiceras spinaries (QUENSTEDT,1858).

Descriptions based on type material:1878 Arietites stellaris SOWERBY. – HERBICH, p. 77, pl.

20 C, fig. 2 a, b.1908 Arietites obtusus SOW. sp. var. vulgaris nov. var. –

VADÁSZ, p. 377, pl. 10, figs. 2-4, 4a, 5, text-fig. 27.

Synonymy:1858 Euagassiceras spinaries. – QUENSTEDT, pl. 11, figs.

8-9.1953 Arietites (Asteroceras) obtusus SOW. var. vulgaris

VADÁSZ. – RAILEANU (in PREDA & RAILEANU), p.350, pl. 9, fig. 2.

1995 Euagassiceras spinaries (QUENSTEDT). – CORNA &DOMMERGUES, p. 34, pl. 3, fig. 4, text-fig. 5 (cumsyn.).

Type material: VADÁSZ (1908) studied several well-preserved specimens of which he selected no holo-type. Several syntypes, including previously figuredspecimens, still exist in the collection.

Measurements:

D U WH WW WW/WHNo. 12.579.H (a) 52.1 21.7 16.5 16.6 1.00No. 12.579.H (b) 41.3 18.0 13.3 15.0 1.12No. 5916.H 43.8 16 – – –

Remarks: The syntypes represent subadult stages withsubquadratic whorl section and characteristic coarse or-

namentation that consists of strong radial ribs that areprojected forward at the well-developed ventro-lateraltubercles. The venter is carinate-bisulcate. This form iscommon in our newly collected material and is nowconfidently identified with Euagassiceras spinaries.

Strat ig raphic dis t r ibut ion: Early Sinemurian, Semi-costatum Zone.

Arietites subrejectus VADÁSZ, 1908Fig. 6e

Revised name: Coroniceras cf. rotiforme (SOWERBY,1824).

Descriptions based on type material:1908 Arietites subrejectus nov. sp. – VADÁSZ, p. 381, pl.

10, fig. 6, text-fig. 28.

Synonymy:1953 Arietites (Arnioceras) subrejectus VADÁSZ. –

RAILEANU (in PREDA & RAILEANU), p. 353.

Type material: Specimen No. 5921.H, an internalmould of phragmocone, is holotype by monotypy ofthis nominal species.

Measurements:


Remarks: It is unjustifiable to maintain this incomplete,poorly preserved specimen as a separate species. Althoughour identification remains tentative, it most likely representsthe inner whorls of Coroniceras rotiforme, mainly on thebasis of its ribbing and the occasional ventro-lateralnodes.

Strat ig raphic dis t r ibut ion: Early Sinemurian.

Arietites pseudospiralis VADÁSZ, 1908Fig. 6c

Revised name: Coroniceras? sp.

Descriptions based on type material:1878 Arietites Conybeari SOWERBY. – HERBICH, p. 106,

pl. 20B, fig. 1a, b.partim 1878 Arietites multicostatus SOWERBY. – HERBICH,

p. 106, pl. 20B, fig. 1a, b.1908 Arietites pseudospiralis nov. sp. – VADÁSZ, p. 382,

pl. 11, fig. 4, text-fig. 29, 29a.

Synonymy:1953 Arietites (Arnioceras) pseudospiralis VADÁSZ. –

RAILEANU (in PREDA & RAILEANU), p. 353, pl. 9,fig. 4.


Type material: Of the two syntypes of VADÁSZ (1908),only No. 5920.H, the original of his text-fig. 29 isfound in the collection, although in a less completestate.

Measurements:

D U WH WW WW/WHSyntype 93.5 70.9 23.7 21.3 0.89

Remarks: VADÁSZ (1908) based this species on two spe-cimens illustrated by HERBICH (1878) (one as conybeari,the other as multicostatus). One of the syntypes is stillavailable, but revision of this form is made difficult by theproblem of homeomorphy in Early Jurassic arietitids. Therelatively sparse, rursiradiate ribs that bear faint ventro-lateral nodes suggest a Coroniceras?, although this form isalso close to some species of Metophioceras.


Arietites carenatus FUCINI var. antiqua VADÁSZ, 1908Fig. 6f

Revised name: Coroniceras cf. rotiforme (SOWERBY,1824).

Descriptions based on type material:1908 Arietites carenatus FUC. sp. var. antiqua nov. var. –

VADÁSZ, p. 384, pl. 10, fig. 7, 7a, b, text-fig. 30.

Synonymy:1953 Arietites (Arnioceras) carenatus FUCINI, cfr. var.

antiqua VADÁSZ – RAILEANU (in PREDA &RAILEANU), p. 353.

Type material: Specimen No. 5919.H, a poorly pre-served, incomplete internal mould, is holotype bymonotypy of this nominal subspecies.

Measurements:


Remarks: This form resembles Arietites subrejectusVADÁSZ, 1908, and it is similarly unjustifiable to maintainthis incomplete specimen as a well-defined subspecies.Although our identification remains tentative, it most likelyrepresents the inner whorls of Coroniceras rotiforme,mainly on the basis of its ribbing and the occasional ventro-lateral nodes.


Arietites spiratissimus QUENSTEDT var. simplexVADÁSZ, 1908

Fig. 6j

Revised name: Metophioceras? simplex (VADÁSZ,1908).

Descriptions based on type material:1908 Arietites spiratissimus QUENST. sp. var. simplex

nov. var. – VADÁSZ, p. 386, pl. 11, fig. 1, 1a, text-fig. 31.

Synonymy: 1953 Arietites (Asperoceras) spiratissimus QUENST. sp.

var. simplex VADÁSZ. – RAILEANU (in PREDA &RAILEANU), p. 350.

Type material: Specimen No. 5927.H, a worn butotherwise well-preserved internal mould, is holotypeby monotypy.

Measurements:


Remarks: An almost complete, evolute specimen of largesize, strong, dense, rursiradiate ribbing, and tricarinate-bisulcate venter that is difficult to identify with otherestablished taxa. VADÁSZ proposed it as a subspecies ofVermiceras spiratissimus, a form with significantly moreevolute coiling. Tentatively we elevate simplex to thespecies rank and assign it to Metophioceras, also notingaffinity to Arnioceras, that cannot be excluded in the lack ofpreservation of innermost whorls.

Arietites ultraspiratum FUCINI var. costosa VADÁSZ,1908

Original descr ipt ion:

1908 Arietites ultraspiratum, FUC. sp. var. costosa, nov.var. – VADÁSZ, p. 387, text-fig. 32.

Type material: No type specimen of this subspecies,which lacks an original illustration by VADÁSZ (1908),has been identified in the collection, therefore it ispresumed lost.

Arietites altesulcatus WÄHNER var. involutaVADÁSZ, 1908

Fig. 6g

Revised name: Caenisites plotti (REYNES, 1879).

Descriptions based on type material:1908 Arietites altesulcatus WÄHN. var. involuta nov. var.

– VADÁSZ, p. 393, pl. 11, fig. 3, text-fig. 34.

Synonymy:1995 Caenisites plotti (REYNES). – CORNA & DOMMER-

GUES, p. 36, pl. 4, fig. 2, text-figs. 6-7 (cum syn.).




Type material: Specimen No. 5924.H, a complete andmoderately well-preserved internal mould, is holotypeby monotypy.

Measurements:


Remarks: This form of moderately rapidly expandingevolute whorls, strongly carinate-bisulcate venter, andslightly adorally arched ribs is best identified withCaenisites plotti as re-described recently by CORNA &DOMMERGUES (1995). A peculiar feature is the irregularityin costation manifest in a looped rib that is preceded by avery short intercalated rib. VADÁSZ (1908) speculates thatthe specimen may be figured as Arietites multicostatusSOWERBY in HERBICH (1878, pl. 20A, fig. 2a-c), but thiscannot be confirmed.

3.5. Species erected by TOULA (1911)

Rhacophyllites lexeni TOULA, 1911Fig. 6i

Revised name: Lytoceras ‘lexeni’ (TOULA, 1911).

Descriptions based on type material:1911 Rhacophyllites Lexeni n. sp. – TOULA, p. 20, pl. 2,

fig. 3.

Type material: TOULA (1911) states that the twostudied specimens were obtained by F. PODEK, aprivate collector from Kronstadt (now Brasov) andsubsequently deposited in the regional Saxon Museum(„Burzenländer Sächsisches Museum“) in Brasov. Thefigured specimen, a fully septate internal mould, hasbeen discovered in the collection of the GeologicalInstitute of Hungary (inventory no. J.11475) and isdesignated here as lectotype.

Measurements:


Remarks: The labels accompanying the specimen suggestthat VADÁSZ, sometime after publication of both TOULA’s(1911) and his main work (1908), revised the originalidentification as Lytoceras secernendum STEF. var. lexeni,although his opinion remained unpublished. The volution,general lack of ornament, and septal sutures suggest thatthis form indeed belongs to Lytoceras. The lack of bodychamber hinders a full characterization of this form. Thespecies name ‘lexeni’ is tentatively retained here, noting thatit appears obsolete and should likely be replaced once amodern revision of several closely similar Lytoceras specieswill be available to decide on its synonymy.

3.6. Species and subspecies erected byRAILEANU (in PREDA & RAILEANU, 1953)

Rhacophyllites (Dasyceras) rakosensis HERBICH

acuticostatus RAILEANU, 1953

Revised name: Juraphyllites nardii (MENEGHINI, 1853).

Descriptions based on type material:1953 Rhacophyllites (Dasyceras) rackosensis (sic!) HERB.

var. acuticostatus (nov. var.). – RAILEANU (in PREDA

& RAILEANU), p. 339, pl. 1, fig. 2, 2a.

Remarks: The subspecies was proposed on the basis ofits strong ribbing on body-chamber and the suture line asdiagnostic features. We cannot confirm the existence ofthe type material. Based on the original description andfigure in PREDA & RAILEANU (1953), the specimen isidentified as Juraphyllites nardii (MENEGHINI); for furtherdiscussion, see taxon Ammonites transylvanicus HAUER,1866.

Rhacophyllites (Paradasyceras) ürmösensisHERBICH tenuilobata RAILEANU, 1953

Revised name: Paradasyceras uermoesense (HERBICH,1878).

Descriptions based on type material:1953 Rhacophyllites (Paradasyceras) ürmösensis HERB.

var. tenuilobata (nov. var.). – RAILEANU (in PREDA

& RAILEANU), p. 340, pl. 3, fig. 1; pl. 5, fig. 2.

Synonymy:1996 Paradasyceras tenuilobata RAILEANU. – POPA &

PATRULIUS, pl. 16, fig. 2.

Remarks: The subspecies was proposed on the basis ofits narrow umbilicus as a diagnostic feature. However,umbilical width varies rather continuously within the rangeof variability of the species, therefore maintaining thesubspecies is not warranted. (See also taxon Phyllocerasürmösense HERBICH, 1878).

Lytoceras elegans RAILEANU, 1953

Descriptions based on type material:1953 Lytoceras elegans nov. sp. – RAILEANU (in PREDA &

RAILEANU), p. 344.

Remarks: The original description is not accompanied byany illustration and the type specimen was not available tostudy, therefore no revision is attempted here.


Schlotheimia boucaultiana D’ORBIGNY ombilicataRAILEANU, 1953

1953 Schlotheimia boucaultiana D’ORB. ombilicata (nov.var.) – RAILEANU (in PREDA & RAILEANU), p. 349.

Remarks: The original description is not accompanied byany illustration and the type specimen was not available tostudy, therefore no revision is attempted here.

Arietites (Vermiceras) juliae BONARELLI

retroversicostatus RAILEANU, 1953

Revised name: Vermiceras? sp.

Descriptions based on type material:1953 Arietites (Vermiceras) juliae BON. retroversico-

status (nov. var.). – RAILEANU (in PREDA &RAILEANU), p. 355, pl. 10, fig. 3.

Remarks: Without examination of the type specimen andonly based on the provided description it’s hard to give anexact determination. The specimen has elliptical whorlcross section, wider than high. The ornamentation consistsof radial ribs and the venter bears a keel sided by two deepsulci. The whorl shape and the ornamentation indicate aVermiceras but identification at the species level is notattempted here.

4. Genera based on type species from thePersani Mts.

4.1. Dasyceras HYATT, 1900

The genus Dasyceras was introduced as a new name,without diagnosis or description, in listing the generaof the family Phylloceratidae (HYATT, 1900). Phyllo-ceras rakosense HERBICH was designated as its typespecies. It is only known from a single specimen, itsholotype appears somewhat abraded that explains itsunusual ornamentation, exaggerated in the illustrationby HERBICH (1978). This form is easily accommo-dated in Juraphyllites (see also discussion of taxonPhylloceras rákosense HERBICH, 1878). Therefore weregard Dasyceras as a junior subjective synonym ofJuraphyllites.

4.2. Schistophylloceras HYATT, 1900

Schistophylloceras was introduced in the same work(HYATT, 1900) and in the same fashion as Dasyceras,with no diagnosis or description, purely by de-signation of Phylloceras aulonotum HERBICH as itstype species. In this case, however, a distinctiveventral sulcus on internal moulds that may correspond

to a keel on shelly specimens represents a unique dia-gnostic feature within the phylloceratids that permitsreliable identification of the genus. See also discusionof taxon Phylloceras aulonotum HERBICH, 1878.

4.3. Paradasyceras SPATH, 1923

The genus Paradasyceras was proposed by SPATH

(1923) to include the group of Rhacophyllites stella(SOWERBY) but Phylloceras uermoesense HERBICH

was designated as its type species. The arising con-fusion is discussed in detail and resolved by RAKÚS

(1993), who introduced the new genus Togaticeras forthe stella group, thereby retaining Paradasyceras as amore clearly established genus of Early Jurassic jura-phyllitids. See also discussion of taxon Phyllocerasürmösense HERBICH, 1878.

4.4. Tragolytoceras SPATH, 1924

SPATH (1924) proposed the genus Tragolytoceras with-out a diagnosis, designating Lytoceras altecinctum(HAUER) as its type species with unambiguousreference to the specimen figured by BONARELLI

(1900). As demonstrated above, HAUER’s taxon (seeAmmonites altecinctus HAUER, 1866) is best regardedas nomen dubium, presumably a subjective juniorsynonym of Ectocentrites petersi. Further confusionarises from SPATH’s disregard of the type specimenand his explicit, albeit invalid, fixation of BONARELLI’sspecimen typifying the genus. BONARLLI’s specimenis herein regarded a separate species, and its originalmisidentification with A. altecinctus HAUER is ex-plained by the limitations imposed by the fragmentarytype of the latter taxon. Clearly, SPATH, whose famili-arity with the type of altecinctus is doubtful, merelyfollowed the misidentification of BONARELLI. In caseof misidentified type species, Article 70.3 of ICZN(1999) allows selection of another type species thatbest serves stability of nomenclature. In a revisionof the genus, WIEDMANN (1970) also interpretedTragolytoceras as typified by the characters ofBONARELLI’s specimen. CORNA et al. (1997) furtherdemonstrates that this practice remains the prevailingusage. Designating BONARELLI’s specimen herein asthe holotype of Tragolytoceras bonarellii nov. sp., andfixing the new species itself the type species of thegenus therefore best preserves stability. In order tovalidate such type fixation, we have referred thiscase to the International Commission on ZoologicalNomenclature.




Tragolytoceras bonarellii nov. sp.Fig. 4c

partim 1900 Lytoceras altecinctum (HAUER). – BONARELLI,p. 70, pl. 9, fig. 1 (non pl. 8, fig. 12).

1957 Tragolytoceras altecinctum (HAUER). – ARKELL, p.L194, text-fig. 224/3.

Etymology: Named after G. BONARELLI, Italianpaleontologist who first described and figuredthe specimen selected for holotype of the newspecies.

Holotype: Specimen No. C.8480, in the MorenaCollection of the Department of Palaeontology, TheNatural History Museum (London).

Measurements:

D U WH WW WW/WHHolotype 46 22 14 8 0.57

Diagnosis : Evolute shell of compressed, mode-rately rapidly enlarging whorls with stages of differentornamentation. Constrictions on early whorls yield to

Table 1. Alphabetical index of ammonoid species and subspecies originally named from the Early Jurassic localitynear Racos (Alsórákos) and their revision based on the studied type material.


gradually developing coarse, blunt, rectiradiate ribsthat cross the venter.

Descr ipt ion: Shell evolute, compressed, discoidal.High ellipsoid whorls of moderately rapid expansion(~1.75); umbilicus wide (48% of diameter). Ornamentchanging, innermost whorls bear widely spaced con-strictions. Coarse, blunt ribbing develops gradually,following a stage with subdued ornament. Last pre-served half whorl bears 13 primary and 5 secondaryribs. Ribs straight, rectiradiate, arise at umbilicalmargin and cross the venter lending a capricornappearance. Irregular intercalatory ribs appear at mid-flank.

Remarks: The holotype was first figured byBONARELLI (pl. 9, fig. 1) and refigured in the Treatise(ARKELL, 1957). The other specimen of BONARELLI

(pl. 8, fig. 12), refigured by WIEDMANN (1970: 996,pl. 9, fig. 3), differs in having sharper, prorsiradiateribs that commonly bifurcate at the umbilical margin,and develop earlier in ontogeny. This form may re-present a different species therefore we chose not toinclude it in the type series.

Occur rence: Foci, Pontalto (or Ponte Alto), Apen-nines, Italy, from Sinemurian strata.

5. Conclusions

The Tepei (Töpe) Creek near Racos (Alsórákos) inthe Persani Mts. is a classical Hettangian–Sinemurianammonoid locality in “Rosso Ammonitico” facies ofthe Tethyan province. Since its discovery in 1866, sixdifferent authors described 37 new nominal species orsubspecies from here. The major part of the type mate-rial exists in the collection of the Babes-Bolyai Uni-versity in Cluj. After a revision, we retain 12 speciesas valid (one of which is elevated from subspeciesrank), and regard 14 nominal taxa as subjective juniorsynonyms of other species. Three subspecies appearunjustified to distinguish within their species. Wedeem the types of three nominal taxa unidentifiable atthe species level therefore recommend abandoningthese names. One species is tentatively retained until arevision of related taxa, whereas the unavailabilityof types precluded the revision of another four taxa.Considering all nominal taxa, seven lectotypes aredesignated herein. Confirming the existence of typematerial suppresses previous selection of neotypes for

three species. Four species has served as type speciesof genera, of which Tragolytoceras was marred byconfusion, necessitating the introduction of a newspecies (T. bonarellii) and its fixation as the new typespecies of the genus. The main results of our revisionare summarized in Table 1.

Acknowledgements

We are grateful to the following colleagues for providingaccess to collections or loan of specimens of their in-stitutions: I. BUCUR, S. FILIPESCU, and M. POPA (Depart-ment of Geology-Paleontology, Babes-Bolyai University,Cluj-Napoca). L. KORDOS (Geological Institute of Hungary,Budapest), S. BAKER (The Natural History Museum,London), L. MARTIRE and G. PAVIA (Department of EarthSciences, University of Torino). I. GYORFFY was instru-mental in initiating this study. I. FOZY (Hungarian NaturalHistory Museum) provided one of the photographs. P. L.SMITH and J. GUEX are thanked for their helpful reviews.Financial support was received through a Bolyai Fellowshipto JP, from the Hungarian Academy of Sciences, and theHungarian Research Fund (OTKA) Grant T042802.

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Manuscript received: September 1st, 2006.Revised version accepted by the Stuttgart editor: December11th, 2006.

Addresses of the authors:

ROBERT TOMAS, Department of Geology-Paleontology,Babes-Bolyai University, Mihail Kogalniceanu nr. 1, Cluj-Napoca, RO - 400084 Romania;Present address: S.C. Deva Gold S.A., Piata Unirii nr. 9,RO-330152, Deva, Jud. Hunedoara, Romania;E-mail: [email protected]

JÓZSEF PÁLFY (corresponding author), Hungarian Academyof Sciences–Hungarian Natural History Museum, ResearchGroup for Paleontology, POB 137, Budapest, H-1431Hungary; E-mail: [email protected]


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