THE SYSTEMATIC POSITION OF PSEUDOERTLISPONGUS LAHM · THE SYSTEMATIC POSITION OF PSEUDOERTLISPONGUS LAHM (RADIOLARIA) AND DESCRIPTION OF SOME NEW MIDDLE TRIASSIC AND LIASSIC RADIOLARIAN

Geol. Paläont. Mitt. Innsbruck, ISSN 0378-6870, Sonderband 4, S. 287-299, 1996

THE SYSTEMATIC POSITION OF PSEUDOERTLISPONGUS LAHM(RADIOLARIA) AND DESCRIPTION OF SOME NEW MIDDLE TRIASSIC

AND LIASSIC RADIOLARIAN TAXA

Heinz W.Kozur

With 1 plate

Abstract:Pseudoertlispongus LAHM is a transitional form between Paroertlispongus KOZUR & MOSTLER and Oertlispongus DUMITRICÄ,

KOZUR & MOSTLER.

Paroertlispongus, Pseudoertlispongus and Oertlispongus appeared in successive order at the base of the Lower Anisian Par-oertlispongus diacanthus Zone, at the base of the upper Subzone {Yeharaia annulata Subzone) of the uppermost Anisian Spon-gosilicarmiger transitus Zone and at the base of the Lower Ladinian Spongosilicarmiger italiens Zone (= base of Reitziitesre/fz/Zones.str.)One new genus, 7 new species and one new subspecies of Middle Triassic radiolarians and the Liassic genus Liassobetracchi-um n. gen. are described.

Zusammenfassung :Pseudoertlispongus LAHM ist eine Übergangsform zwischen Paroertlispongus KOZUR & MOSTLER und Oertlispongus DUMI-

TRICÄ, KOZUR & MOSTLER. Paroertlispongus, Pseudoertlispongus und Oertlispongus erscheinen in stratigraphischer Abfolgean der Basis der unteranisischen Paroertlispongus diacanthus Zone, in der oberen Subzone {Yeharaia annulata Subzone) derhoch-oberanisischen Spongosilicarmiger transitus Zone und an der Basis der unterladinischen Spongosilicarmiger italicusZone (= Basis der Reitziites reitziiZone).Eine neue Gattung, 7 neue Arten und eine neue Unterart mitteltriassi sehen Radiolarien die liassische Gattung Liassobetracchi-um werden beschrieben.

1. Introduction

KOZUR & MOSTLER ( 1994) regarded Pseudoert-lispongus LAHM, 1984 as a junior synonym ofPa-roertlispongus. At that time, only one species ofPseudoertlispongus was known that occurred inthe middle Fassanian together with advanced Pa-roertlispongus KOZUR & MOSTLER, advancedOertlispongus DUMITRICÄ, KOZUR & MOSTLER,

Baumgartneria DUMITRICÄ and Falcispongus DU-

MITRICÄ. Shell sculpture and arrangement of thespines on the shell of Pseudoertlispongus corre-spond to Paroertlispongus and also the roundcross section of the main polar spine throughout itslength is typical for Paroertlispongus. The only

difference is a slightly curved posterior end of themain polar spine. This last feature is transitional toOertlispongus that has a recurved main polar spinewith a short straight part and a long recurved part.However, the previously known distribution ofPseudoertlispongus with first appearance after thefirst appearance of Oertlispongus did not favourthe possibility that Pseudoertlispongus is a transi-tional form between Paroertlispongus and Oert-lispongus.

Now we have found rich Pseudoertlispongusfaunas before the first appearance of Oertlispon-gus in Hungary as well as in Sicily. Therefore, theformerly known occurrence of Pseudoertlispon-gus weddigei was only the uppermost range of

287

Pseudoertlispongus together with advanced Oert-lispongus that ranges, however, up to the Longo-bardian. The first appearance of Paroertlispongus,Pseudoertlispongus and Oertlispongus is therefo-re in clear stratigraphie order, independently fromthe fact that the primitive Oertlispongidae co-occur with advanced ones. The oldest known oc-currence of Paroertlispongus is in the upperLower Anisian Paroertlispongus diacanthusZone. There, only forms with long, slender, intheir middle part not widened main polar spineoccur, and the opposite polar spine is not much dif-ferent. Advanced Paroertlispongus with wideningof the main polar spine in or after its mid-lengthappear in the Upper Anisian Tetraspinocyrtis lae-vis Zone from the Paraceratites trinodosus Zoneof the ammonoid zonation. They ranges up to theupper Fassanian.

Pseudoertlispongus with terminally slightlycurved main polar spine appeared in the upper-most Anisian upper Subzone (Yeharaia annulataSubzone) of the Spongosilicarmiger transitusZone. With very slight changes, this genus rangesup to the upper Fassanian.

Oertlispongus with recurved main polar spinebegins at the base of the Ladinian Spongosilicar-miger italicus Zone (base of the Reitziites reitzizone s.str. in the ammonoid zonation). The oldestOertlispongus species, O. primus n. sp., is a primi-tive form with a long straight part of the main polarspine that is only a little shorter than or as long asthe curved part that is only very little recurved. Thebent is already slightly flattened (oval cross sec-tion) as typical for several Oertlispongus species,but never present in Paroertlispongus and Pseud-oertlispongus. This genus underwent a rapid evo-lution and advanced forms with very short straightpart of the main spine occur from the middle Fas-sanian up to the Middle Longobardian. Contem-poraneous with O. primus, a closely related formappeared, in which, however, the inward curvedpart is short, the bent of the main spine is stronglyflattened and on its outer side slightly widened.These forms represent the oldest Falcispongus, F.zapfei n. sp., which is the ancestral form of the ad-vanced oertlispongids with strongly differentiatedmain polar spine that characterize the Tethyan and

partly also the Circum-Pacific Ladinian and Cor-devolian.

In the present paper, the stratigraphically veryimportant species from the transitional field of Par-oertlispongus, Oertlispongus and Falcispongusare described. Furthermore some other new spe-cies and genera are described that are used in stra-tigraphical papers without systematic part, toavoid the use of nomina nuda.

2. Systematic part

Subclass Radiolaria MÜLLER, 1858Order Polycystida EHRENBERG, 1838

Suborder Entactinaria KOZUR & MOSTLER, 1982Superfamily Palaeoscenidiacea RIEDEL, 1967

emend. KOZUR & MOSTLER, 1982Family EptingiidaeDuMiTRicÄ, 1978

Genus Cryptostephanidium DUMITRICÄ, 1978

Type species: Cryptostephanidium cornigerumDUMITRICÄ, 1978

Cryptostephanidium sugiyamai n. sp.

Deri vatio nominis: In honour of Prof. Dr. KAZU-

HIRO SUGIYAMA, Nagoya, for his excellent work onLower and Middle Triassic radiolarians.Holotypus: The specimen figured by SUGIYAMA

(1992), Fig. 13-3; rep.-no. ESN 146172.Locus typicus: Mt. Kinkazan section of Gifu,Gifu Prefecture, central Japan.Stratum typicum: Bedded cherts, sample KIN61 (see SUGIYAMA, 1992, Fig. 2), Lower AnisianHozmadia gifuensis Zone.Diagnosis: Pore frames with verrucose to irregu-lar spiny surface. The three main spines are of une-qual length. Their proximal third to half is three-bladed, their distal half is round.Description: Shell small, subspherical. The surfa-ce of the pore frames is verrucose or covered withvery small, irregular, partly oblique spines. Poresmoderately large, mostly pentagonal. The three

288 Geol. Paläont. Mitt. Innsbruck, Sonderband 4, 1996

main spines are situated in continuation of spines ofthe entactinarian spicular system. Their widthdecreases gradually from the base to the pointedneedle-like distal end. The spine Ais longer than thetwo spines L. the proximal third to half of the mainspines is three-bladed. The furrows between theblades are wide, at the base of the spines deep, butthey become shallow in distal direction. The distalhalf to 2/3 of the main spines has a circular crosssection and no blades and furrows are present.Measurements:Diameter of shell: 85-100 urnLength of main spines: 60-95 urnMaximal width of the main spines: 20-25 urnOccurrence: Common in the Lower Anisian of Ka-raburun peninsula (western Turkey) and in Japan.Remarks: The Turkish material is rich in speci-mens, but not well preserved. Therefore, the speci-men figured by SUGIYAMA (1992) was chosen asholotype.

Cryptostephanidium sugiyamai n. sp. is a tran-sitional form between Spongostephanidium Du-MITRICÄ, 1978 emend. KOZUR, KRAINER & MOST-

LER, this volume, and Cryptostephanidium DUMI-

TRICÄ, 1978. The proximal third to half of the mainspines is already incannate as in Cryptostepha-nidium, the distal half to 2/3 of the main spines isneedle-like with round cross section as in Spongo-stephanidium.

In the younger Cryptostephanidium speciesnearly the entire main spines are tricarinate. In theotherwise similar C. verrucosum DUMTTRICÄ,

1978, the three main spines have equal length.

Suborder Spumellaria EHRENBERG, 1875Superfamily Actinommacea HAECKEL, 1862

emend. KOZUR & MOSTLER, 1979Family Pantanelliidae PESSAGNO, 1977

Genus Liassobetraccium n. gen.

Type species: Betraccium bavaricum KOZUR &MOSTLER, 1990

Deri vatio nominis: According the similarity toBetraccium PESSAGNO and the occurrence in theLiassic.

Diagnosis: Cortical shell subspherical with highand coarse pore frames. The pores are very large,but irregular in size and shape. Most pores are tri-angular and tetragonal, but some pentagonal poresmay be also present. Some pores may be dividedby fragile secondary pore frames. Vertices of thepore frames with distinct nodes and spines. Thethree stout, three-bladed main spines are twistedor untwisted, equidistant and of the same size. Me-dullary shell present.Occurrence: Hettangian of theTethys.Assigned species:Betraccium bavaricum KOZUR & MOSTLER, 1990Betraccium hettangicum KOZUR & MOSTLER,

1990Betraccium inaequiporatum KOZUR & MOSTLER,

1990Betraccium verticispinosum KOZUR & MOSTLER,

1990Remarks: Betraccium PESSAGNO, 1979, is a verycommon Upper Norian and Rhaetian genus bothin the Tethys and in the Circum-Pacific realm. Itsuddenly disappeared at the base of the Liassic(CARTER, 1993). However, there are some closelyrelated species in the Hettangian that have been as-signed to Betraccium by KOZUR & MOSTLER

(1990). In contrast to typical Triassic Betracciumspecies, in all these species the pore frames enclo-ses pores of different size and shape. In Betracci-um all pores have the same size and pentagonal orhexagonal outline. In the Liassic forms the poreoutline is in general triangular or tetragonal andonly subordinately pentagonal outline of the poresis present. These Liassic species group is separa-ted as Liassobetraccium n. gen.

Superfamily Sponguracea HAECKEL, 1862emend. KOZUR & MOSTLER, 1981

Family Oertlispongidae KOZUR & MOSTLER,

1980 (in DUMITRICÄ et al., 1980)

Genus Falcispongus DUMITRICÄ, 1982

Type species: Falcispongus falciformis DUMITRI-

CÄ, 1982

Geol. Paläont. Mitt. Innsbruck, Sonderband 4, 1996 289

Falcispongus zapf ei n. sp.(PL 1,Fig. 10)

Derivatio nominis: in honour of Prof. emer. H.ZAPFE, Vienna, for his outstanding contributionsto the Triassic stratigraphy.Holotypus: The specimen on PI. 1, Fig. 10; rep.-no. 14-4-94/V-5Locus typicus: Outcrop ca. 350 m south of Pietradei Saracini, Sosio Valley, western Sicily (Italy).Stratum typicum: Basal 20 cm of red, strongly sili-ceous, marly limestones, sample Ko 5. Very baseof the Ladinian, base of the S. italiens Zone.Material: 2 specimens.Diagnosis: The perpendicularly inward bent di-stal part of the main polar spine is considerablyshorter than straight axial proximal part. Mainpolar spine at the bent flattened, with small wide-ning at the outer side directly before the bent.Description: Straight axial proximal part of themain polar spine in its upper part flat, with smallwidening at its outer side directly before the bent.The perpendicularly inward bent distal part is di-stinctly shorter than the straight part of the mainpolar spine. It is not recurved. Shell not preserved.Measurements:Length of the straight part of the main polar spine:300-360 umWidth of the straight part of the main polar spine atits proximal end (above the conical part that wasoriginally within the shell): 30-33 umWidth of the straight part of the main polar spinedirectly before the bent: 40-53 urnLength of the inward curved part of the main polarspine: 160-200 urnOccurrence: Lower part of Oertlispongus primi-tivus Subzone of the S. itálicas Zone of western Si-cily (lower Fassanian).Remarks: Falcispongus zapfei n. sp. is the mostprimitive Falcispongus species. All other lowerLadinian species of this genus have a long inwardcurved and mostly also recurved distal part that islonger than the straight proximal part. The wide-ning at the outer side of the distal end of thestraight part of the main polar spine or (and) at thebent is more pronounced in the other Falcispon-gus species.

F zapfei is similar both to Pseudoertlispongusmostleri siciliensis n. subsp. and to Oertlispongusprimus n. sp. The main polar spine of Pseudoert-lispongus mostleri siciliensis is not flattened nearthe bent and no widening at the outer side of thestraight part of the main polar spine directly belowthe bent is present. In Oertlispongus primus n. sp.,the inward curved part is about as long as thestraight part. Apparently, at the base of the Ladini-an both the first primitive Oertlispongus and thefirst primitive Falcispongus have evolved fromPseudoertlispongus of the P. mostleri group.

Genus OertlispongusDUMITRICÄ, KOZUR & MOSTLER, 1980

Type species: Oertlispongus inaequispinosus Du-MITRICÄ, KOZUR & MOSTLER, 1980

Oertlispongus primus n. sp.(PI. 1, Fig. 9)

Derivatio nominis: Oldest known true Oertli-spongus species.Holotype: The specimen on PI. 1, Fig. 9; rep.-no.14-4-94/V-24.Locus typicus: Outcrop ca. 350 m south of Pietradei Saracini, Sosio Valley, western Sicily (Italy).Stratum typicum: Basal 20 cm of red, stronglysiliceous, marly liMestones, sample Ko 5. Verybase of the Ladinian, base of the S. italicus Zone.Material: 5 specimens.Diagnosis: Straight proximal part of the mainpolar spine long, only a little shorter than or as longas the nearly perpendicularly inward curved partthat is only insignificantly recurved.Description: The straight axial proximal part ofthe main polar spine is long for the genus and onlysomewhat shorter than the inward curved part. Inthe proximal part the cross section of the mainpolar spine is round, but in its upper part and in thebent the main polar spine is flattened and has anoval cross section. The inward curved part liesnearly perpendicular to the straight axial part; it isonly somewhat longer than or as long as the axial


straight proximal part. The inward curved part isnearly not recurved. Shell not preserved.Measurements:Length of the straight axial proximal part of themain polar spine: 280-295 urnWidth of the straight axial proximal part of themain polar spine at its proximal end (above the co-nical part that was originally within the shell):27-29 urnWidth of the main polar spine directly before thebent: 39^43 urnLength of the inward curved part of the main polarspine: 280-305 urnOccurrence: Lower part of Oertlispongus primi-tivas Subzone of the S. italiens Zone. Basal part ofLadinian. Hungary and Sicily (Italy).Remarks: Oertlispongus primus n. sp. is the ol-dest and most primitive Oertlispongus species. Ithas transitional character to Pseudoertlispongusmostleri siciliensis. It is distinguished from allPseudoertlispongus species by its longer, stronglyinward curved part that is nevertheless not or onlya little longer than the straight axial proximal partof the main polar spine. Additionally, the mainpolar spine is flattened near the bent.

In all other Oertlispongus species the inwardbent part is considerably longer than the straightproximal part and, except of few undescribedforms from the lowermost Ladinian, strongly re-curved.

Falcispongus nicorae n. sp. displays a shorterinward bent part and a slight widening is present atthe outer side of the bent.

Genus ParoertlispongusKOZUR & MOSTLER, 1981

Type species: Paroertlispongus multispinosusKOZUR & MOSTLER, 1981

Remarks: Three Paroertlispongus species occurin sample Ko 5 from the basal S. italiens Zone ofSosio Valley, Sicily (Italy), that are P. multispino-sus KOZUR & MOSTLER, 1981 (PL 1, Fig. 1), P. ra-

rispinosus KOZUR & MOSTLER, 1981 (PL 1, Fig. 2)and P. chinensis (FENG, 1992) (PL 1, Fig. 3).Thefirst two species are only represented by isolated

main polar spines, a very typical preservation forOertlispongidae. The main polar spine of P. multi-spinosus is widest after its midlength. The mainpolar spine of P. rarispinosus is cylindrical in itsproximal half and becomes needle-like towards itsdistal end. P. chinensis has several small, low ribsat the base of both polar spines.

Genus Pseudoertlispongus LAHM, 1984

Type species: Pseudoertlispongus weddigeiLAHM, 1984

Occurrence: Yeharaia annnlata Subzone of theSpongosilicarmiger transitus Zone (upper Illyr-ian)oftheTethys.Remarks: Pseudoertlispongus has the same dis-tribution of the second order spines on the spongyshell as Paroertlispongus, but the distal end of themain polar spine is curved. The straight part of themain polar spine is considerably longer than thecurved part. In Oertlispongus the distribution ofthe second order spines is different from Paroert-lispongus and Pseudoertlispongus, and the mainspine is recurved. The straight part of the mainpolar spine is shorter than the curved part. By thepresence of a distally somewhat curved spine,Pseudoertlispongus is a transitional form betweenParoertlispongus and Oertlispongus. It appearsconsiderably later than the first Paroertlispongus,but distinctly before the first Oertlispongus.

Pseudoertlispongus angulatus n. sp.(PL 1, Figs. 4,5)

Deri vatio nominis: According to the abruptly in-ward curved distal end of the main polar spine.Holotypus: The specimen on PL 1, Fig. 4; rep.-no.16-2-95/H-104.Locus typicus: Torrente San Calogero section(see CATALANO, DI STEFANO & KOZUR, 1991).Stratum typicum: Basal 10 cm of the gray radio-larite directly above the Changxingian red deep-water clay. Uppermost Anisian Y annulata Subzo-ne of the S. transitus Zone.Material: 12 specimens.


Diagnosis: The main polar spine is widest imme-diately before the abrupt, angular bent in its distalthird.Description: The main polar spine is large. Itswidth increases gradually and slowly until thesharp angular bent in the distal third of the mainpolar spine. The straight, inward directed distalpart of the main polar spine has different length andthe angle with the straight axis of the main polarspine varies, but it is not larger than 45°. Only thisinward directed part becomes narrower towardsthe pointed distal end. Shell not preserved.Measurements:Length of the main polar spine: 440-510 jamLength of the straight axial part of the main polarspine: 350-370Length of the inward bent part: 100-150 urnWidth of the proximal part of the main polar spine(above the conical part that was originally situatedinside the shell): 30 urnWidth directly before the bent of the main polarspine: 35^6 jamOccurrence: Uppermost Anisian (? and lower-most Ladinian) of western Sicily.Remarks: In the other two known Pseudoertlis-pongus species, P. weddigei LAHM, 1984 and P.mostleri n. sp., the bent is gently curved. In P. wed-digei additionally the main spine becomes narro-wer directly before the bent.

Pseudoertlispongus mostleri n. sp.(PI. 1, Figs. 6,7)

1994 Paroertlispongus weddigei (LAHM, 1994)- KOZUR & MOSTLER, p. 69-70, PL 12,Figs. 12-14

Deri vatio nominis: In honour of Univ.-Prof. Dr.H. MOSTLER for his outstanding contributions tothe Triassic radiolarian research.Holotypus: The specimen on PL 1, Fig. 6; 14-4-94/V-3.Locus typicus: Outcrop ca. 350 m south of Pietradei Saracini, Sosio Valley, western Sicily (Italy).Stratum typicum: Basal 20 cm of red, strongly sili-ceous, marly limestones, sample Ko 5. Very baseof the Ladinian, base of the S. italiens Zone.

Material: 35 specimens.Diagnosis: Shell spongy with numerous by-spi-nes that are not concentrated to any part of theshell. Main polar spine cylindrical, widest directlybefore or at the bent. The bent is gradual androunded.Description: The spherical spongy shell consistsof several layers. Its surface is covered by nume-rous needle-like by-spines that start in well preser-ved forms from node-like elevations. The mainpolar spine is very long and mostly the only pre-served part of a specimen. It is cylindrical and wi-dest directly before the bent or at the bent. The bentis gradually rounded. The inward-curved distalpart of the main polar spine is short, its angle withthe long axial straight part of the main polar spineis around 45°, but in the subspecies P. mostleri si-ciliensis between 45° and nearly 90°. The secondpolar spine opposite or somewhat obliquely oppo-site to the main polar spine is long, needle-like, butrarely preserved.Measurements: See under the subspecies.Occurrence: Uppermost Anisian Y. annulataSubzone of the S. transitus Zone and lower Ladi-nian S. italiens Zone.Remarks: Pseudoertlispongus mostleri is the tran-sition form between Pseudoertlispongus andOertlispongus. The spongy shell with nodes andby-spines on the entire surface corresponds to Pa-roertlispongus multispinosus. The main polarspine is also similar, but distally inward curved.

In Oertlispongus primus n. sp., the oldest Oert-lispongus species, the early perpendicularly in-ward curved part is longer than or as long as thestraight axial part of the main polar spine.

Pseudoertlispongus mostleri mostleri n. subsp.(PL 1, Fig. 6)

Holotypus, locus typicus and stratum typicum:As for the species.Material: 3 \ specimens.Diagnosis: With the character of the species. Thedistal, inward bent part of the main polar spine isalways short and its angle with the axial straightpart of the main polar spine is not larger than 45°.


The main polar spine is broadest directly beforethe bent.Measurements:Diameter of shell: 180-267 urnLength of the main polar spine: 350-430 jimWidth of the main polar spine at its proximal end(outside the shell or outside the conical part, if theshell is not preserved: 20-35 urnWidth of the main polar spine directly before thebent: 26-50 urnLength of the inward curved distal part of the mainpolar spine: 50-95 urnLength of the by-spines: 10-50 |umOccurrence: Uppermost Anisian Y annulataSubzone of the S. transitus Zone and Lower Fassa-nian S. italicus Zone of Hungary, Southern Alpsand Sicily (Italy).Remarks: In Pseudoertlispongus mostleri sicili-ensis n. subsp., the main polar spine is widest at thebent and the distal part is stronger inward curved.

Pseudoertlispongus mostleri siciliensis n. subsp.(PI. 1, Fig. 7)

Derivatio nominis: According to the occurrencein Sicily.Holotype: The specimen on PI. 1, Fig. 7; rep.-no.:14-4-94/V-2.Locus typicus and stratum typicum: As for thespecies.Material: 4 specimens.Diagnosis: With the character of the species.Main polar spine widest at the bent or of the samewidth throughout its straight axial part. Inwardbent strong, partly nearly perpendicular to thestraight axial part of the main spine.Measurements:Length of the main polar spine: 500-520 urnWidth of the main polar spine at its proximal end(after the conical part, that was originally insidethe shell): 40-47 urnWidth of the main polar spine at the bent: 43-47 urnLength of the inward curved distal part óf the mainpolar spine: 133-147 urnOccurrence: Upper part of the uppermost Anisi-an Y annulata Subzone of the S. transitus zone and

lower part of Oertlispongusprimitivus Subzone ofthe S. italicus Zone.Remarks: Pseudoertlispongus mostleri mostlerin. subsp. has a fewer inward bent distal part of themain polar spine that is widest directly before thebent.

In Oertlispongus primus n. sp. the inward partof the main polar spine is somewhat longer than oras long as the straight axial part of the main polarspine.

Superfamily Trematodiscacea HAECKEL, 1862emend. KOZUR & MOSTLER, 1979

Family Relindellidae KOZUR & MOSTLER, 1980(inDuMTTRicÄetal., 1980)

Genus Pentaspongodiscus KOZUR & MOSTLER,

1979

Type species: Pentaspongodiscus tortilis KOZUR

& MOSTLER, 1979

Pentaspongodiscus lahmi n. sp.

1984 Pentaspongodiscus anisicum KOZUR &MOSTLER 1981 -LAHM, p. 57, PI. 10, Fig. 1

Derivatio nominis: In honour of Dr. B. LAHM,

München, who firstly found first this species.Holotypus: The specimen figured by LAHM (1984)on PI. 10, Fig. 1 under Pentaspongodiscus anisi-cus KOZUR & MOSTLER, 1981 ; rep.-no. Prot. 3623.Locus typicus: Section Ree. A, 2.5 km SE of Re-coaro, in the Val di Creme.Stratum typicum: Limestone with siliceousnodules about 40 cm above the beginning of thepelagic limestones, sample Ree. A 9. Lower S. ita-licus Zone (Lower Ladinian).Material: 23 specimens.Diagnosis: The five untwisted tricarinate mainspines are slender and shorter than the shell diame-ter.Description: The discoidal shell is large compa-red with the length of the main spines. Its equatori-al outline is round. It is spongy with Alievum pat-tern of the outer pore frames. Distinct small nodes


are present on the slightly elevated vertices. Fivefragile short rays originate in all vertices. The fiveslender main spines are tricarinate, with a needle-like, round distal prolongation. They are untwi-sted and always shorter than the shell diameter.Measurements:Diameter of shells: 150-167 umLength of the main spines : 100-13 3 urnWidth of the main spines at their base: 23-25 urnOccurrence: Lower Ladinian S. italicus Zone ofHungary and Italy.Remarks: LAHM (1984) assigned this species toPentaspongodiscus anisicus KOZUR & MOSTLER,

1981. However, this Upper Anisian species (ance-stor of P. lahmi) has a pentagonal equatorial outli-ne of the shell and the robust, long main spines aredistinctly longer than the shell diameter.

Suborder Nassellaria EHRENBERG, 1875Family Hinedorcidae KOZUR & MOSTLER, 1981

Genus Picapora KOZUR & MOSTLER, 1981

Type species: Picapora robusta KOZUR & MOST-

LER, 1981

Picaporafassanica n. sp.

Derivatio nominis: According to the occurrencein the Fassanian.Holotypus: The specimen figured by GORICAN &BusER(1990),Pl.ll,Fig. 1.Locus typicus: 1.5 km north of Zaklanec nearHorjul at the Zaklanec-Koreno road (southern partof the Slovenian Basin); see GORICAN & BUSER,

1990, Fig. 1.Stratum typicum: Cherty limestone below vol-canics, sample Gr 10 (see GORICAN & BUSER,

1990, Fig. 2).Material: 23 specimens.Diagnosis: APicapora with small vertical horn.Description: Cephalis subspherical (somewhatbroader than high) with small pores that are mostlyclosed by a layer of microgranular silica; surfaceslightly verrucose. The apical horn is stout, tricari-

nate, with deep and broad furrows between thehigh and narrow blades. A distinct, but rather smalland short, carínate vertical horn is present. Thoraxlarger than cephalis, with the same surface structu-re and mostly closed small pores. It has the shape ofa cone frustum or it is subspherical. Its wide apertu-re is closed by a fragile thin layer with small poresthat is often destroyed. The three feet are stout, mo-derately long, tricarinate. They run from the distalpart of the cephalis on the surface of the thorax andare free only beyond the thorax. Cephalic spicularsystem with Mb, A, V, 2 L, D, 21. The spines A andV continue outside the shell as apical horn and hornV, whereas D and 2 L continue as feet.Measurements:Height of cephalis: 30-40 urnWidth of cephalis: 50-55 urnHeight of thorax: 50-60 urnWidth of thorax: 70-85 urnLength of the apical horn: 40-45 mLength of the free part of the feet: 50-55 urnOccurrence: Fassanian of Slovenia, Hungary andItaly.Remarks: The Julian Picapora robusta KOZUR &MOSTLER, 1981 has a very big vertical horn.

Family TriassocampidaéKOZUR & MOSTLER, 1981

Genus Praeyeharaia n. gen.

Type species: Yeharaia transita KOZUR & MOST-

LER, 1994Derivatio nominis: According to the ancestralposition to Yeharaia NAKASEKO & NISHIMURA,

1979Diagnosis: Test multicyrtid, conical. Cephalis co-nical, smooth with small apical horn. Thorax a litt-le broader, but in general shorter than the cephalis,ring-like or hoop-like, with nodes or a ring ofnodes; pores small, mainly closed. Abdomen cy-lindrical or hoop-like, nodose, with small, partlyopen pores. The postabdominal segments are ingeneral inverted conical with a smooth, in the pro-ximal segments slightly nodose proximal ring.The distal part of these segments is nodose. Below


the proximal ring a ring of rather large pores is pre-sent. In the distal part of the postabdominal seg-ments a second, often incomplete pore ring is pre-sent. The proximal two postabdominal segmentsmay be hoop-like, with indistinct proximal ring.Cephalic spicular system with A, V, 2 L, D, 2 1.Only spine A continues outside the shell wall asshort apical spine. Sometimes also a tiny spine ispresent in prolongation of D.Occurrence: Uppermost Anisian S. transitasZone and Lower Ladinian S. itálicas Zone.Assigned species:Yeharaia transita KOZUR & MOSTLER, 1994Yeharaia japónica NAKASEKO & NISHIMURA,

1979Yeharaia lata KOZUR & MOSTLER, 1994Remarks: Yeharaia NAKASEKO & NISHIMURA,

1979, has a long apical spine and an inflated thoraxand the postabdominal segments (except the firstone) have only one ring of pores. TriassocampeDUMITRICÄ, KOZUR & MOSTLER, 1980 has no api-

cal spine.Praeyeharaia n. gen. is transitional between

Yeharaia and Triassocampe. Praeyeharaia spe-cies have been in previous paper partly assigned toYeharaia, partly to Triassocampe.

3. Stratigraphie evaluation

The Oertlispongidae are the most important La-dinian radiolarian guideforms of the Tethys. Inprevious studies, Paroertlispongus was found inthe Illyrian and Fassanian, and rich Oertlispongusfaunas with O. inaequispinosus primitivus KOZUR

& MOSTLER were found in the lower Fassanianlower Reitziites reitzi ammonoid zone s. str. (O. in-aequispinosus primitivus Subzone of the S. itali-ens radiolarian zone). Despite the fact that Paro-ertlispongus was regarded as the forerunner ofOertlispongus (KOZUR & MOSTLER, 1981), thetransition forms between both genera, and the im-mediate forerunner of Falcispongus were notfound.

In the upper subzone of the S. italicus Zone bothadvanced Oertlispongus with very short proximal

straight part of the main polar spine (e.g. O. inae-quispinosus inaequispinosus) and typical Falcis-pongus were found.

Investigations of rich further material of sam-ple 100 D (Kellneritesfelsoeoersensis ammonoidzone) from the Felsóórs section in the BalatonHighland (Hungary) and investigations near thebase of the Reitziites reitzi Zone in the BalatonHighland, Southern Alps and in Sicily have yiel-ded the complete transitional series from Paroert-lispongus to Oertlispongus. Pseudoertlispongusmostleri, n.sp. a transitional form between Par-oertlispongus and Oertlispongus appeared nearthe base of the Kellneritesfelsoeoersensis ammo-noid zone. The radiolarian fauna of this level be-longs to the upper subzone (Yeharaia annulataSubzone) of the S. transitus Zone. Beside of Ladi-nian elements, as Eptingium manfredi manfrediDUMITRICÄ, Pseudoertlispongus mostleri n. sp.,Triassocampe deweveri (NAKASEKO & NISHIMU-

RA), T scalaris DUMITRICÄ, KOZUR & MOSTLER

and Yeharaia annulata NAKASEKO & NISHIMURA,

this fauna contains still dominating Anisian ele-ments, listed in KOZUR, KRAINER & MOSTLER

(this volume). Consequently, this fauna was as-signed by KOZUR (1995) to the uppermost Anisi-an. It has, however, transitional character to Ladi-nian radiolarian faunas, but advanced Oertlispon-gidae, which dominate in the entire Ladinian, arestill missing. They begin near the base of the Reit-ziites reitzi ammonoid zone at the base of the S.italicus radiolarian zone. There, not only Pseud-oertlispongus mostleri n.sp. is present, but alsothe first Oertlispongus s. str. in which the lateralcurved (and mainly distinctly recurved) distal partof the main polar spine is at least as long as the pro-ximal straight part. In typical Oertlispongus, thisdistal part is strongly recurved and considerablylonger than the straight proximal part. In Oertlis-pongus primus the inward curved part is about aslong as the proximal straight part and it is therefo-re a perfect transition form between Pseudoertlis-pongus mostleri siciliensis and Oertlispongus in-aequispinosus. In the same level also the first Fal-cispongus, F. zapf ei, appeared. This form also de-rived from Pseudoertlispongus mostleri sicilien-sis, the inward curved part is. still as short as in this


species and considerably shorter than the straightproximal part of the main polar spine, but it is al-ready strongly flattened near the bent, and direct-ly below the bent a small widening is present atthe outer side of the straight part of the main polarspine. By this can be demonstrated that Falcis-pongus did not evolve from Oertlispongus, butdirectly from Pseudoertlispongus.

Throughout the Ladinian, the primitive Oert-lispongidae continue beside the advanced forms.Pseudoertlispongus mostleri n.sp. and P. angula-tus n.sp. are the only Oertlispongidae with diffe-rentiated main polar spine (inward curved distalpart) in the uppermost Illyrian Y. annulata Subzo-ne of the S. transitus Zone. Advanced Oertlispon-gidae with short straight proximal part of the mainpolar spine and long, recurved distal part are notyet present. These forms appeared at the base ofthe S. italicus Zone. In a very short interval theseadvanced forms are only represented by transitio-nal forms to the primitive Oertlispongidae withvery long straight proximal part of the main polarspine, but a little later already forms with verylong recurved distal part and distinctly shortstraight proximal part of the main polar spine arepresent. However, the primitive Oertlispongidaestill continue in this stratigraphie level, e.g. Paro-ertlispongus, Pseudoertlispongus and primitiveOertlispongus. In the Lower Longobardian high-ly advanced Oertlispongidae with strongly diffe-rentiated spines occur (genera Pterospongus,Bogdanella etc.), but O. inaequispinosus from theFassanian continued into this level. Therefore, forthe subdivision of the Middle Triassic radiolarianfaunas, the first appearance of a certain taxon isvery important and the exact age is indicated bythe most advanced forms within a phylogeneticcline.

Acknowledgements

The author is very grateful for the financial sup-port by the Deutsche Forschungsgemeinschaftand for the support by Univ. Prof. Dr. H. MOSTLER,

Innsbruck.

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Author's address:Dr. sc. H.einz W. Kozur, Rézsü u. 83, H-1029Budapest, Hungary.


Plate 1

If not otherwise indicated, all figured specimens are from sample Ko 5 (basal 20 cm of red, strongly siliceous, marly lime-stones; very base of the Ladinian, base of the S. itálicas Zone) in an Scythian to Ladinian section ca. 350 m south of Pietra deiSaracini, Sosio Valley, western Sicily, Italy (location see GULLO & KOZUR, 1993).

Fig. 1: Paroeertlispongus multispinosus KOZUR&MOSTLER, 1981, isolated main polar spine, x 130, rep.-no. 14-4-94/V-10.

Fig. 2: Paroertlispongus rarispinosus KOZUR &MOSTLER, 1981, isolated main polar spine, X 150, rep.-no. 14-4-94/V-7

Fig. 3 : Paroertlispongus chinensis (FENG, 1992), x 100, rep.-no. 14-4-94/V-8

Figs. 4,5: Pseudoertlispongus angulatus n.sp., x'200, sample SCK11/93, lowermost 10 cm of gray radiolarite directly aboveChangxingian red deep-water clay, Yeharaia annulata Subzone of Spongosilicarmiger transitus Zone, uppermostAnisian, Torrente San Calogero section (see CATALANO et al., 1991); Fig. 4: holotype, rep.-no. 16-2-95/II-104;Fig. 5 : rep.-no. 16-2-95/II-106. .

Fig. 6: Pseudoertlispongus mostleri mostleri n. sp., holotype, x 200, rep.-no. 14-4-94/V-3.Fig. 7: Pseudoertlispongus mostleri siciliensis n. subsp., holotype, x 200, rep.-no. 14-4-94/V-2.

Fig. 8: EptingiummanfredimanfrediDVMiTRicÂ., 1978, x 130, rep.-no. 14-4-94/V-9.Fig. 9: Oertlispongus primus n. sp., holotype, x 145, rep.-no. 14-4-94/V-24.

Fig. 10 Falcispongus zapfei n. sp., holotype, x 150, rep.-no. 14-4-94/V-5.Fig. 11 : Triassocampe scalaris baloghi KOZUR & MOSTLER, 1994, x 300, rep.-no. 14-4-94/V-6.


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THE SYSTEMATIC POSITION OF PSEUDOERTLISPONGUS LAHM · THE SYSTEMATIC POSITION OF PSEUDOERTLISPONGUS LAHM (RADIOLARIA) AND DESCRIPTION OF SOME NEW MIDDLE TRIASSIC AND LIASSIC RADIOLARIAN

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