Reichenbacher, Bettina / Gaudant, Jean. Eclogae Geologicae Helvetiae. Band (Jahr): 96 (2003) Heft 3. PDF erstellt am:

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1 On Prolebias eyeri (Agassiz) (Teleoistei, Cyprinodontifores) fro the OligoMiocene of the Upper Rhinegraben area, with the establishent of a new genus and a new species Autor(en): Objekttyp: Reichenbacher, Bettina / Gaudant, Jean Article Zeitschrift: Eclogae Geologicae Helvetiae Band (Jahr): 96 (2003) Heft 3 PDF erstellt a: Persistenter Link: Nutzungsbedingungen Die ETHBibliothek ist Anbieterin der digitalisierten Zeitschriften. Sie besitzt keine Urheberrechte an den Inhalten der Zeitschriften. Die Rechte liegen in der Regel bei den Herausgebern. Die auf der Plattfor eperiodica veröffentlichten Dokuente stehen für nichtkoerzielle Zwecke in Lehre und Forschung sowie für die private Nutzung frei zur Verfügung. Einzelne Dateien oder Ausdrucke aus diese Angebot können zusaen it diesen Nutzungsbedingungen und den korrekten Herkunftsbezeichnungen weitergegeben werden. Das Veröffentlichen von Bildern in Print und OnlinePublikationen ist nur it vorheriger Genehigung der Rechteinhaber erlaubt. Die systeatische Speicherung von Teilen des elektronischen Angebots auf anderen Servern bedarf ebenfalls des schriftlichen Einverständnisses der Rechteinhaber. Haftungsausschluss Alle Angaben erfolgen ohne Gewähr für Vollständigkeit oder Richtigkeit. Es wird keine Haftung übernoen für Schäden durch die Verwendung von Inforationen aus diese OnlineAngebot oder durch das Fehlen von Inforationen. Dies gilt auch für Inhalte Dritter, die über dieses Angebot zugänglich sind. Ein Dienst der ETHBibliothek ETH Zürich, Räistrasse 101, 8092 Zürich, Schweiz,

2 /03/ DOI /S X Birkhäuser Verlag, Basel, 2003 Eclogae geol. Helv. 96 (2003) On Prolebias eyeri (Agassiz) (Teleostei, Cyprinodontifores) fro the OligoMiocene of the Upper Rhinegraben area, with the establishent of a new genus and a new species Bettina Reichenbacher1 & Jean Gaudant2 Key words: Aphanolebias n. gen.. Prolebias. Aphanius. Cyprinodontifores. Oligocene. Miocene ABSTRACT The study of previously and newly collected skeletons wilh otoliths preserved in siili of the cyprinodontifor species Prolebias eyeri (Agassiz) fro the Upper Hydrobia Beds of Frankfurt/Main (Early Miocene. Hanau Basin) enables us to propose an eended diagnosis of this species, which becoes the typespecies of the new genus Aphanolebias. Aphanolebias nov. gen. differs fro Prolebias Sauvage due to its triangularshaped otoliths and fro Aphanius Nardo due to its conical jaw teeth and the shape of its dentary. A younger synony of Aphanolebias eyeri (Agassiz) is Aphanius geraniae Weiler In addition, we have reexained skeletons and otoliths of a Prolebias species fro the Upper Oligocene and Lower Miocene of the Upper Rhine graben and the Mainz and Hanau Basins, which was also assigned to P. eyeri. We introduce the new species P. inalzi for it. We conclude that all otoliths de terined as P. eyeri sensu Weiler (1942: 1963) belong to P. alzi n. sp. Slight differences in otolith orphology ay occur between P. /««/./populations of different ages. RESUME L'étude de spéciens portant des otolithes in situ, à la fois anciens et nouvel leent récoltés, de l'espèce de Cyprinodontifores Prolebias eyeri (Agas siz). du Miocène inférieur de Francfort (Bassin de Hanau. Alleagne), nous conduit à en proposer une diagnose éendée pour cette espèce qui devient l'espècetype du nouveau genre Aphanolebias. Aphanolebias nov. gen. diffère de Prolebias Sauvage par la fore triangulaire de ses otolithes et d'aphanius Nardo notaent par la possession de dents coniques sur les âchoires et par la fore du dentaire. Il est égaleent établi que toutes les otolithes isolées désignées depuis Weiler (1963) coe Aphanius geraniae Weiler doivent être rapportées à l'espèce Aphanolebias eyeri (Agassiz). On a égaleent réexainé les squelettes et les otolithes d'un atériel de véritables Prolebias de l'oligocène supérieur et du Miocène inférieur du Fossé rhénan et des bassins de Mayence et de Hanau, qui avaient été décrits par er reur coe P. eyeri (Agassiz). La nouvelle espèce P. alzi n. sp. est créée pour ce atériel. Toutes les otolithes déterinées depuis Weiler (1942: 1963) coe P. eyeri (Agassiz) sont considérées coe appartenant réelleent à l'espèce P. alzi n. sp. En outre, après avoir discuté la variabilité des oto lithes de cette espèce, il est constaté que de légères différences peuvent exister entre des populations d'âge légèreent différent. ZUSAMMENFASSUNG Prolebias eyeri (Agassiz) (Cyprinodontifores) aus den Oberen HydrobienSchichten von Frankfurt (UnterMiozän. Hanauer Becken) wird anhand von Skeletten it in situ erhaltenen Otolithen untersucht. Das Material stat aus alten und neuen Aufsalungen. Wir geben eine erweiterte Dia gnose dieser Art und definieren sie als TypusArt der neuen Gattung Aphano lebias. Aphanolebias nov. gen. ist von Prolebias Sauvage aufgrund seiner dreieckigen Otolithen und von Aphanius Nardo aufgrund seiner konischen Kieferzähne und der For des Dentale unterschieden. Ein jüngeres Synony von Aphanolebias eyeri (Agassiz) ist Aphanius geraniae Weiler. Eine andere Art aus de OberOligozän und UnterMiozän des Ober rheingrabens, des Mainzer und des Flanauer Beckens, die bisher für Prolebias eyeri gehalten worden war. wird als P. alzi neu definiert. Alle i Sinne von Weiler (1942: 1963) als P. eyeri bestite Otolithen gehören zu P. alzi n. sp. Zwischen P. a/?/populationen unterschiedlicher stratigraphi scher Herkunft sind kleine Unterschiede in der Otolithenorphologie zu ver zeichnen. 1. Introduction and historical review w Malz (1978:, 444) correctly recognized that the sedients The species Lebias eyeri was originally described by yielding Lebias eyeri belong to the Upper Hydrobia Beds Agassiz (1839) fro the Miocene of Frankfurt (Hanau that can be placed in the late Early Miocene (Martini 1987; Basin, Gerany). It based on abundant skeletal aterial Reichenbacher 2000). that was collected between 10 and 16 depth when dig However, it was rather difficult to get an accurate underging the well of the central ceetery (von Meyer 1834). standing of Lebias eyeri Agassiz because its original de Departent of Earth and Environental Sciences & GeoBioCenter. LudwigMaxiilansUniversität Munich. RichardWagnerStr. 10. D80333 Munich. Eail: b.reichenbacher@lrz.uniuenchen.de rue du Docteur Magnan. F75013 Paris. Eail: gaudantfecipgp.jussieu.fr On Prolebias eyeri (Agassiz) fro the OligoMiocene of the Upper Rhinegraben 509

3 / hz K zlz Qr a Kaiserslautern i</ T "* Landa *< Wiesbaden a Mainz fi AINZ^ \ Alzey /Wors UzjYY Mannhei? Hanau FRANKFURT a Darstadt 1 =c Heidelberg DUDENHOFEN <c Speyer1^ / a Wiesloch V SPOCK Q Bruchsal tr*. U Karlsruhe ^ 5 a Rastatt 30 k *: Localities yielding Prolebias alzi n. sp. A Senckenberg locality, yielding Aphanolebias eyeri (AG.) Fig. 1. Map showing the origin of the studied aterials of Aphanolebias ey eri (Agassiz) and Prolebias alzi n.sp. (ap fro Weiler odified). 1112). According to Weiler 1942). the skeletons and otoliths fro the borehole Baden 4 originate fro the "lower Hydro bia Beds" and the "boundary Corbicula/Hydrobia Beds". However, today it is well known that in the Rhinegraben area these strata truly belong to the Upper Cerithiu Beds (Upper Oligocene to Lower Miocene) and to the underlying Middle Cerithiu Beds (Upper Oligocene) (e.g., Reichenbacher 2000). Unfortunately, the aterial fro the borehole Baden 4 cannot be located. Weiler (1963) referred to the figured otolith fro the bore hole Baden 4 as "otolithholotype of Prolebias eyeri". In the sae article, Weiler (1963: 25. pl. 2) described and figured skeletons fro the Middle Cerithiu Beds of the borehole Dudenhofen ( ; Fig. 1) in the southern Rhine graben as P. eyeri. He entioned two skeletons with in situ preserved otoliths and considered that they are siilar to his "otolithholotype of P. eyeri". Additionally, Weiler (1963: Fig. 7679) described and figured isolated otoliths as P. eyeri. However, only the two otoliths fro the Upper Hydrobia Beds ofthe borehole Spöck ( ; Fig. 1) belong to this species (Reichenbacher 2000: 73). Reichenbacher & Mödden (1996), Reichenbacher (2000). and Reichenbacher & Sienknecht (2001) deterined and figured isolated otoliths fro several outcrops and boreholes in the Mainz and Hanau Basins (Fig. 1) as P. eyeri, referring to the deterinations of Weiler ( ). These otoliths originate fro the Landschneckenkalk and the overlying Upper Ceriihiu Beds and Riissingen Foration (forer In flata Beds, Schäfer & Kadolsky 2002). They are absent in the overlying WiesbadenForation (forer Lower Hydrobia Beds, Reichenbacher & Keller 2002), but appear again in the overlying Upper Hydrobia Beds and Landschneckenergel. In the present study we deonstrate on basis of skeletons and otoliths that P. eyeri sensu Weiler (1942, 1963) does not agree with P. eyeri sensu Agassiz (1839). scription is incoplete and soewhat iprecise. Additionally, the original aterial, which belonged to von Meyer's private collection, was lost since that tie, including the two syntypes figured by Agassiz (1839. V, Tab. 41, Figs. 78). According to Sauvage (1874), Lebias eyeri should be in cluded in the fossil genus Prolebias Sauvage It is worth to ention that the one "P. eyeri" skeleton, which is en tioned out ofthe Rhinegraben region (d'eraso 1929: Messin ian fro Senigallia, Italy), probably is a young specien of Aphanius crassicaudus (Agassiz). Weiler (1929) discussed and copared the skeletal charac ters of P. eyeri when introducing the Oligocene species Pro lebias praecursor Weiler, but his coparison based only on the incoplete description by Agassiz (1839). Weiler (1942) reported skeletons of P. eyeri fro the borehole Baden 4 in the southern Upper Rhinegraben. He did neither describe nor illustrate the, but noted that several skeletons were bearing roundshaped otoliths in situ, of which he described and figured one (Weiler 1942: 24, PI. 1, Figs. 2. Material and origin of aterial The studied aterial coes fro previous and new collections and is kept in the sections Palaeozoology I (SMF P) and Mi cropalaeontology (SMF PO) of the Senckenberg Museu Frankfurt. Previous collection of P. eyeri sensu Agassiz 1839) This aterial was collected by H. von Meyer and probably originates fro the Upper Hydrobia Beds. SMF P. 1687b: skeleton fro Frankfurt. Alte Gasse: Fig SMF P. 1686b: skeleton, probably fro sae locality as SMF P. 1687b: Fig SMF P. 1686a: otolith, found in situ in the counterpart of SMF P. 1686b: entioned by Malz (1978): Fig New collection of P. eyeri sensu Agassiz 1839) This aterial originates fro the Upper Hydrobia Beds of Frankfurt. Senckenberganlage. and was collected by the aateur collectors K. Weiss. W. Ott. and M. Keller (Ott 2003). The fossiliferous layers were accessible during the 510 B. Reichenbacher & J. Gaudant

4 Tab. 1. Main characters of Aphanolebias eyeri (Agasse 1839). All skeletons were found in the Upper Flydrobia Beds fro the underground of Frankfurt/iVlain. standard length (SL) previous collection new collection SMFP. 1686ab. SMFP b up to 40 axiu height 1/4 of SL 1/4 to 1/3.5 of SL of body oral jaws preaxilla with preaxilla with conical teeth conical teeth otolith triangular triangular total nuber of vertebrae nuber of 15 (15)16(17) postabdoinal vertebrae shape of caudal fin convex convex nuber of 16 (15)16 principal rays of caudal fin nuber of dorsal fin rays nuber of 910 pterygiophores of dorsal fin nuber of anal fin rays nuber of pterygiophores of anal fin position of anal slightly in front of alost opposed to fin origin of dorsal fin dorsal fin insertion of slightly nearer to slightly nearer to pelvic fins anal fin than to anal fin than to pectoral fins pectoral fins nuber of 1215 pectoral fin rays nuber of 67 pelvic fin rays construction of the new etropolitan line U 4. uniting the central railway sta tion to Bockenheier Warte. Besides P. eyeri. soe gobiid skeletons and one atherinid skeleton were found. According to its otoliths preserved in situ. the latter belongs to Heitrichas bartensteini (Malz). Gobiids do not occur in strata older than the Upper Hydrobia Beds and H. bartensteini is restricted to the Upper Hydrobia Beds (Reichenbacher 2000). Thus, these taxa support the lithostratigraphic position of the fossiliferous layers. SMF P. 9612: skeleton (collected by W. Ott under the nuber Se 26); Fig SMF P. 9613: skeleton (collected by K. Weiss under the nuber Se 2W4): Figs SMF P. 9614: skeleton (collected by W. Ott under the nuber Se 2a4): Figs SMF P. 9615: skeleton (collected by K. Weiss under the nuber Se 2W1); Fig SMF P. 9616: skeleton (collected by W. Ott under the nuber Se 211 ): Fig SMF P : seven skeletons (collected by K. Weiss under the nubers Se2W2. W3.W610). SMF P SMF PO : skeleton with both otoliths in situ (collected by M. Keller): Fig SMF PO : otoliths found in situ (collected by M. Keller). Collection of isolated otoliths of Aphanius geraniae Weiler 1963 SMF P sagittae fro the Upper Hydrobia Beds of Frankfurt/Main. borehole Mainzer Landstraße 148 (22.9 depth) (entioned in Malz 1978). SMF P sagittae fro the Upper Hydrobia Beds of Frankfurt/Main. borehole Mainzer Landstraße 156 (8 depth) (entioned in Malz 1978). Further 45(1 sagittae are kept in the Senckenberg Museu (aterial of Malz 1978). Collection of Prolebias eyeri sensu Weiler ) A. Fro Middle CerithiuBeds of the borehole Dudenhofen 1 ( depth) SMF P. 3509: skeleton: figured in Weiler (1963: PI. 2. Fig. 5); holotype of P. inalzi n. sp.: Fig SMF P. 3366AB: skeleton: figured in Weiler (1963: Figs. 74ab); paratype of P. alzi n. sp.: Figs SMF P. 3572: skeleton: entioned in Weiler (1963): Fig SMF P. 3574: skeleton: entioned in Weiler (1963). SMF P. 9617: skeleton: Fig SMF P. 9618a: skeleton and otolith ebedded in the sedient: Fig. 9. SMF P. 9618bc: four skeletons. B. Fro boreholes and outcrops in the Mainz and Hanau Basins, fro bore holes in the Upper Rhinegraben About 650 isolated otoliths (aterial of Reichenbacher & Mödden 1996; Rei chenbacher 2000). 3. Reexaination ot Prolebias eyeri sensu Agassiz (1839) Previous collection (Figs Tab. 1) Specien SMF P. 1687b (Fig. 2.2) is characterized by its elon gate body, the axiu height of which equals slightly less than 1/4 of the standard length. Its rather short vertebral col un consists of 28 or 29 vertebrae, 15 being postabdoinal. Its caudal fin exhibits a convex posterior outline. The dorsal fin is opposed to the anal fin; it has about 10 rays. The anal fin be gins slightly in front of the dorsal fin; its 12 rays are supported by about 10 pterygiophores. The pelvic fins are located slightly nearer to the origin of the anal fin than to the pectoral fins base. Specien SMF P. 1686b (Fig. 2.1) provided additional inforation by showing soe conical teeth in the preaxilla. Malz (1978) isolated one in situ preserved otolith (Fig. 3.1) in the counterpart of SMF P. 1686b. It is a right sagitta exhibit ing a syetrical triangular shape and a rather proinent ros tru, slightly corroded at its tip. The sulcus, situated in the iddle of the sagitta, displays a sall and narrow ostiu and a long cauda. which is strongly bent down and pointed at its dis tal end. New collection (Figs Tab. 1) These skeletons (Fig. 2.3, ) correspond well with the skeletons fro the previous collection and reveal soe addi On Prolebias eyeri (Agassiz) fro the OligoMiocene of the Upper Rhinegraben 511

5 1 Lz''* c r ZgpfjV ' i. a 7 C':l / fe^ T» «s* " *«v v&* :*t WgßiE^ "Sr ' M / *c _,'. * T fctf. $4.r,. jfjä:. W,J fs r,/ w* ". 's*...v" M?*»»»*. N ' v. / 5 iwtiï % 'i Ti Wh y//iï &..w$m ss ust ' TT Äst*.,*î*j % ä/zza* ^*j%m> :. Lj^f 5T s^ ff zlé: WtS&Zrtf* M. 5 ' * v. / «! ' nm.. * : é ^ ' n_» : 3****^ ^l H ^**]» v SPS STS*» * *. ^* '*»& ~#î tyf,s^~ ""S *» v,; 5 ÜsTj Fig. 2. Aphanolebias eyeri (AOASSiz). General view of body of three speciens. Photographs D. Serrette Specien SMF Specien SMF Specien SMF Upper Hydrobia Beds, probably Alte Gasse. Frankfurt/Main. Upper Hydrobia Beds. Alte Gasse. Frankfurt/Main Upper Hydrobia Beds. Senckenberganlage. Frankfurt/Main. P. 1686b. P. 1687b. P. tional inforations. They deonstrate jaws provided with any sall conical teeth (Fig. 4) distributed in several rows, a dentary which is devoid of any spurlike edial process (Fig. 5). a relatively high nuber of principal caudal finrays (Figs ) and two hypural plates incopletely fused cauda bent down and pointed. A slight variability occurs rela tively to the shape and length of the rostru. (Fig. 6.4). The studied skeletons of P. eyeri sensu Agassiz (1839) dis play skeletal and dental characters that are well known fro fossil Prolebias species (e.g.. Gaudant : Gaudant & Reichenbacher 2002). However, it sees that not Soe skeletons revealed in situ preserved otoliths (Figs ). These otoliths (sagittae) are triangular with a proi nent rostru and display a sulcus with the distal end of the 512 B. Reichenbacher & J. Gaudant 4. Discussion and introduction ot Aphanolebias nov. gen.

6 Mx Pap"' * Px I 1 I ^ Fig. 4. Aphanolebias eyeri (Agassiz). Lateral view of the right upper jaw of specien SMF P fro the Upper Hydrobia Beds. Senckenberganlage. Frankfurt Main. Mx: axilla: Px: preaxilla exhibiting lwi7conic.il teeth. Fig. 3. Aphanolebias eyeri (Agassiz). Each scale bar B. Reichenbacher Photographs 13. Otoliths preserved in situ in skeletons. 1. Specien SMF P. 1686a. right sagitta fro the counterpart of specien SMF P. 1686b (see Fig. 2.1). Upper Hydrobia Beds, probably Alte Gasse. Frankfurt/Main. 2. Left sagitta fro specien Se 214 of W. Ott's private collection. Upper Hydrobia Beds. Senckenberganlage, Frankfurt/Main. 3. Specien SMF PO Left (3a) and right (3b) sagitla fro specien SMF P Upper Hydrobia Beds. Senckenberganlage. Frankfurt/Main. 47. Isolated otoliths, previously deterined as Aphanius geraniae Weiler. 45. Speciens SMF P left sagittae fro the Upper Hydrobia Beds of Frankfurt/Main, borehole Mainzer Landstraße 148 (22.9 depth) (fro Malz 1978). 67.Speciens SMF P. 5554ab. righi sagittae fro the Upper Hydrobia Beds of Frankfurt/Main, borehole Mainzer Landstraße 156 (8 depth) (fro Malz 1978). >v, 1 ':_ Fig. 5. Aphanolebias eyeri (Agassiz). Inner view of the right dentary of specien SMF P fro the Upper Hydrobia Beds. Senckenberganlage. Frankfurt/Main. lx any derived skeletal or dental character appears in Prolebias. Consequently, no definition of Prolebias fro a phylogenetic point of view was given so far (Sethi 1960; Parenti 1981). We consider the following characters of Prolebias Sauvage as ost significant. Altogether, they allow to distin guish Prolebias fro Aphanius Nardo and Valencia Myers. jaws provided with conical teeth (tricuspid in Aphanius. conical in Valencia) dentary without any spurlike edial process (with process in Aphanius. without in Valencia) otolith with a dorsal tip and rounded (without dorsal tip and triangular in Aphanius and Valencia) It should be added that the typical otoliths of Prolebias (with dorsal tip and rounded) have been found in several species in situ. e.g.. in P. hungaricus Gaudant 1991, in P. cephalotes (Agassiz) (unpublished data), and in P. aff. weileri (Salis) (Gaudant & Reichenbacher 2002). The triangularshaped otoliths of recent Aphanius species were illustrated by Malz (1978) and Reichenbacher & Sienknecht (2001). Valencia otoliths, which are also triangularshaped, are described and illustrated in Sienknecht & Reichenbacher (in prep.). It should also be added that Valencia is not known as fossil, whereas Prolebias is only known as fossil. Aphanius is known as fossil since the Early Miocene (Burdigalian, 1718 Ma; Gau dant & RoviraSendrós 1998). Fro our description of P. eyeri sensu Agassiz (1839) it is evident that this species bears characters that neither allow to assign it to Prolebias Sauvage, nor to include it in the genus Aphanius Nardo or Valencia Myers. Its general osteology, the conical teeth of its jaws, and the shape of the dentary are evidently typical for Prolebias. But contrary to Prolebias. its otoliths are triangularshaped and reseble recent Aphanius and Valencia otoliths. Consequently, it is necessary to create a new genus for taking into account this unique cobination. On Prolebias eyeri (Agassiz) fro the OligoMiocene of the Upper Rhinegraben 513

7 HHr.* A rt _ <t IV is; g " r :' f.1 e "JJ % _x 2 r * ' T4> *> «2 r... vas SW s ' '_ &J** i Ì * < ra /A tttrr* V ^C. «r./» s.** z I %SS 52 ass Z ;«*" && tz: &C ùtr s '.: MrttMÏ.*,*.' \%J i ji. >. : y Mt ^ :.'f. 1 2 g 2 Fig. 6. Aphanolebias eyeri (Agassiz). Upper Hy drobia Beds. Senckenberganlage. Frankfurt Main. Photographs D. Serrette. 1. Head of specien SMF P Lower jaw of specien SMF P Middle part of body of specien SMF P Caudal axial skeleton of specien SMF P B. Reichenbacher & J. Gaudant

8 Tab. 2. Coparison of Aphanolebias eyeri (Agassiz). Prolebias alzi n. sp.. P. praecursor WEILER, and P. rhenanus Gaudant. Aphanolebias eyeri Prolebias alzi n. sp. Prolebias praecursor Prolebias rhenanus (Agassiz) [= P. eyeri sensu Weiler 1929 Gaudant 1981 (see Tab. 1 Weiler ) Heitrichas) Standard length (SL) up to 40 up to 29 about 40 up to 27 axiu height of body 1/4 to 1/3.5 of SL 1/4 to 1/4.5 of SL not known 1/3 to 1/4.5 of SL jaws preaxilla with conical teeth not known noi known not known otolith triangular rounded not known not known total nuber of vertebrae nuber of postabdoinal (15)16(17) vertebrae shape of caudal fin convex truncated indented truncated nuber of principal rays of (15) caudal fin nuber of dorsal fin rays 912 (79 branched) nuber of pterygiophores not known 910 of dorsal fin nuber of anal fin rays 1114(911 branched) nuber of pterygiophores not known 1112 of anal fin position of anal fin alost exactly significantly in front in front of origin of slightly in front of opposed to dorsal fin of origin of dorsal fin dorsal fin origin of dorsal fin insertion of pelvic fins slightly nearer to anal nearer to pectoral nearer to anal fin very near to anal fin fin than to pectoral fins fins than to anal fin than to pectoral fins nuber of pectoral fin rays not known 1314 nuber of pelvic fin rays 67 6, soeties with clawlike distal ends not known 6 Aphanolebias nov. gen. Derivatio noinis Copound nae ephasizing the unique cobination of characters. Type species Lebias eyeri Agassiz Synonyy of the type species Prolebias eyeri (Agassiz) (not P. eyeri sensu Weiler 1942, 1963, see below), Aphanius geraniae Weiler. Type locality Frankfurt/Main (Hanau Basin, Gerany). Other localities Boreholes in the Upper Rhinegraben (as A. geraniae Weiler, Reichenbacher 2000: Fig. 33). Type foration Upper Hydrobia Beds. Other forations Landschneckenergel, which overly the Upper Hydrobia Beds. Age and stratigraphic range Late Early Miocene (Burdigalian). probably 1920 Ma (Martini 1987, Reichenbacher 2000). Diagnosis Skeletal characters like those of Prolebias Sauvage, but otoliths are triangularshaped and siilar to those of Apha nius Nardo and Valencia Myers. Description ofthe type species (Table 1) Sall cyprinodontifor. the standard length does not ex ceed 40. Body elongated, its axiu height equals 1/5 to 1/3.5 of standard length. Vertebral colun consists of about 2830 vertebrae, 1516 postabdoinal. Caudal fin rounded, with 1516 principal rays. Dorsal fin consisting of iii+i+79 rays supported by 910 pterygiophores. Anal fin beginning ore or less exactly opposed to dorsal fin and consisting of iii+i+911 rays supported by 1012 pterygio phores. Antedorsal and anteanal distance generally ranging fro 69 to 72% of standard length. Pelvic fins situated nearer to the origin of the anal fin than to the base of the pectorals. Dentary and preaxilla provided with sall con ical teeth distributed in several rows. Otoliths (sagitta) tri On Prolebias eyeri (Agassiz) fro the OligoMiocene of the Upper Rhinegraben 515

9 'ä'<#9&sb f. r ri «Liie *~v V., OT,'.n. j.',', rf Zs. 5^ > tri Fk YW Z >C T'Cj W. ff Ì.' Str.. :,,.,, Zi T^.!*# ",,.. fi?.».*<> ita fe.*; 4 ff'f itsxs. se i&. >* *ZV.. r»? C. S: rff fi.. '.i *y> i: Z M.1 <.. an t. Y & à*. &>^3?. œ. ~r.7». >> ê ^' jr.: '*»>sv ' : s,* 'PZ >* V" r? ** ft : Y.i_i Ms 2 l?^ as i'è'j' "V? C»» " _*7 *_...,«Ts, 'zl '.:; f: W* ** t'z» Y! 3M x e'efffis*..r...i'f.fy" tiri «&#&$&.!<rf. ^LnM^YF^. : 7Ì.3 *> >.. rf7! J<r r. ^zzyg e % if 75 _, : ', ""*.» 1. Specien SMF Specien SMF P. f > S*. %*%&fr. a..a. :.. :'j.< Mtr:< «««P isspiyârc: J fr S*. ^.tfrzzjg, '.w: 5 i (southern Upper Rhinegraben). Photographs D. Serrette. 3366A. paratype. holotype. P. 3509, angularshaped with a proinent rostru and a cauda tinctly bent down and pointed at its distal end. dis Synonyies The otoliths oi Aphanolebias eyeri (Agassiz) correspond perfectly with Aphanius geraniae Weiler 1963 (Figs ). which is an otolith basedspecies. 5. Reexaination of Prolebias eyeri sensu Weiler 1942,1963) 5.7 Skeletons and otoliths fro the boreholes Dudenhofen and Baden 4 I Skeletons and otoliths fro the Middle and Upper Cerithiu Beds of the boreholes Dudenhofen and Baden 4 (southern Upper Rhinegraben) were deterined by Weiler (1942, 1963) as Prolebias eyeri (Agassiz). However, this aterial reveals skeletal characters and otoliths, which are clearly different fro Prolebias eyeri (Agassiz) (Tab. 2) JZx^f, i~>~~ i... Fig. 7. Prolebias inalzi n. sp. Middle Cerithiu Beds (Upper Oligocene), borehole Dudenhofen 2. /JC f.fi«% Y Y.'.' J..I '."âjb* '. : *rrji'jrjsl ssj &v_ffï%. 36* L»& B. Reichenbacher & J. Gaudant The skeletons fro the borehole Dudenhofen (Figs. 78) have a standard length ranging fro 13.5 to 29. Their ax iu height of body equals 1/4 to 1/4.5 of standard length. The head (Fig. 8.1) exhibits characters that are well known for the genus Prolebias Sauvage, especially a dentary without spurlike process. The vertebral colun consists of about 30 vertebrae, 1819 being postabdoinal. The caudal fin is poste riorly truncated and includes 1012 principal rays (articulated and furcated) and. both ventrally and dorsally, 78 procurrent rays. It is supported by a caudal axial skeleton in which two distinct hypural plates are present (Fig. 8.4). The dorsal fin consists of iii+i+7 (rarely 8) rays, its principal rays are artic ulated and furcated. It is supported by 7 or 8 pterygiophores (Figs ). The anal fin begins rather significantly in front of the origin of the dorsal fin (Figs. 8.2, 8.3); the anteanal dis tance generally ranges fro 59 to 62.5% of standard length, whereas the antedorsal distance ranges fro 62 to 63.5%. The anal fin consists of ii+i+1114, its principal rays are articulated 1

10 «^ t*'xxr ff *1F'"Z 4'r.Z' ' WM Sì,.'î3ffi r.: i re" z z 0G jv :, a ;.*: a bf/rr*» ft y iffig :'i r 3»* >*' 'f. 30 *T^' wmyéêê feoi* H A.*< Çp^PJfl.rZf&ÜZr : Cf 7LÄ ** tiw r 2 aj:^ SA * >* r** à z <%$? ZZ*. 'issgg ***. ' Yr^s,<V *.jj>>f $3 2 V T fe*^^ rf rfì. *: cr?z Y.Ktn ut ;v, '.>. *S..s,_.. t j *M Y. wss a_*^'/^~ 2 r ^^^t^*^ r>w «r» nl,..tr^f'/ lê^^zxyizrpzc U4 *. ';K { fr; c *r«*«ä<?sr. *&*_{_ '.4 s».v >.' f*"7* ~ s, ft >" TOM* K^lgpy* ' ** <ÄRf i«fig. 8. Prolebias inalzi n. sp. Middle Cerithiu Beds (Upper Oligocene), borehole Dudenhofen 1. (southern Upper Rhinegraben). Photographs D. Serrette * i: V 1 4. Anterior part of body of specien SMF P Middle part of body of specien SMF P. 3366B. Middle part of body of specien SMF P Caudal axial skeleton of specien SMF P. 3366B. On Prolebias eyeri (Agassiz) fro the OligoMiocene of the Upper Rhinegraben 517

11 (SMF Fig. 9. Left sagitta of P. alzi n. sp.. ebedded in ihe sedient of the core of the borehole Duden 1 hofen P. 9618a). Photo graph B. Reichenbacher. and furcated. It is supported by 1315 pterygiophores. Ihe three anteriorost pterygiophores the first one being archshaped take place in front of the heapophyse of the first postabdoinal vertebra. The pectoral fins have about ten rays: the distal end of the longest reaches the base of the pelvic fins (Fig. 8.1). The pelvic fins are obviously situated nearer to the base of the pectorals than to the origin of the anal fin (Fig. 8.2). Six rays ake up the pelvic fins. In soe fishes, the pelvic rays are not articulated and exhibit clawlike distal ends, as shown by specien SMF P. 3366A. The body is covered with cycloid scales, the surface of which bears concentric circuii. Weiler (1963: Fig. 75) entioned and figured rather cor roded otoliths preserved in situ in the skeletons. According to hi. these otoliths have a round shape, a broadly rounded, short rostru, and a pointed antirostru. Unfortunately, we found the strongly corroded and not suited for any figure or ore detailed description. However, we found a slightly flattened, but wellpreserved otolith (sagitta) in the core aterial, ebedded in the sedi ent with the inner face down (Fig. 9). The shape of this otolith is round with a dorsal ri showing a slight edian tip. The posterior ri. well rounded, presents a slight edge in the iddle. The rostru is oderately rounded and shows a slight tip in its upper part. It is not longer than the pointed an tirostru. Length of otolith is 0.88 and height is This otolith corresponds well with the socalled otolithholo type of P. eyeri sensu Weiler (1942) fro the borehole Baden 4. On the other hand, it sees to be soewhat ore rounded in shape than the otolith found in the skeleton SMF P in situ (Weiler 1963: Fig. 75), but this ay result fro the bad preservation of the latter. Taking also into account that no other fish species occurs in the cores of the borehole Dudenhofen 1 ( ). we assue that our otolith belongs to the Prolebias species described above. 5.2 Isolated otoliths fro the Mainz und Hanau Basins Isolated otoliths revealing a rounded shape and a slight edi an tip. as described above, are well known fro Upper Oligocene and Lower Miocene sedients of the Mainz and Hanau Basins. A few speciens were also found in the Upper Rhinegraben (Weiler 1963). Referring to Weiler ( ). all these otoliths were de terined as P. eyeri (Agassiz) in subsequent publications (e.g., Wiesner 1967: Best 1975: Reichenbacher & Weidann 1992; Reichenbacher & Mödden 1996: Reichenbacher 2000). However, on basis of their rounded shape it is clear that they cannot belong to /'. eyeri (Agassiz). because that species, which is now Aphanolebias eyeri. has triangularshaped otoliths (Fig. 3). 6. Discussion and definition of Prolebias inalzi n. sp. Fro the preceding descriptions, it clearly appears thai Pro lebias eyeri sensu Weiler ( ) differs Iro Aphanolebias eyeri (Agassiz) fro the Upper Hvdrobia Beds. As it can be seen fro Tab. 2. Aphanolebias escti has less postabdoinal vertebrae, a saller anal fin having only 1114 rays and 1012 pterygiophores. pelvic fins situated slightly nearer to the origin of the anal fin than to the base of the pectoral fins, and triangularshaped otoliths. Consequently, the skeletons and otoliths, so far deterined as Prolebias ey eri sensu Weiler ( ). belong to a new species. Pro lebias alzi n. sp. Prolebias alzt n. sp. Figs. 79 Derivatio noinis This species is dedicated to Dr. Heinz Malz (Brasche), who was the first to question Weiler's deterination of this aterial (Malz 1978:455). Holotype Specien SMF P (Fig. 7.2), already figured by Weiler (1963. PI. 2. Fig. 5). Type locality Borehole Dudenhofen 1 ( ). southern Upper Rhinegraben (Fig. 1). Other localities Boreholes Grünstadt and Spöck 1 in the Upper Rhine graben. boreholes and outcrops in the Mainz and Hanau Basins (as P. eyeri sensu Weiler ; e.g.. Rei chenbacher 2000: 73). Type foration Middle Cerithiu Beds. Other forations Landschneckenkalk. LJpper Cerithiu Beds. Rüssingen Foration (forer Inflata Beds). Upper Hydrobia Beds. Landschneckenergel (Reichenbacher 2000: 73). Age and chronostratigraphic range Upper Oligocene ( Ma) until Late Early Miocene (1920 Ma). Definition and description Body elongate with a standard length up to 30. a axi u height, which equals 1/4 to 1/4.5 of standard length, and coprising about 30 vertebrae being postab doinal. The caudal fin is truncated and consists of B. Reichenbacher & J. Gaudant

12 principal rays. The dorsal fin is opposed to the anal fin and consists of 78 rays supported by 78 pterygiophores. The anal fin. which begins rather significantly in front of the dorsal fin. displays 1417 rays supported by 1315 ptery giophores. The pectoral fins consist of 10 rays and are of oderate size. The pelvic fins are situated nearer to the pectoral fins than to the origin of the anal fin: they consist of 6 rays having soeties clawlike extreities. Body covered with cycloid scales. Otoliths of rounded shape with,i slili dors.il up and a rostru, which is not or onh slightly longer than the antirostru. Coparison with other species The holotype and only known specien of Prolebias praecur sor Wl n i R 1929 was found in Upper Oligocene sedients near Alzey in the Mainz Basin, either in the Cyrena Marls, or the socalled.süßwasserschichten, or the Upper Ceriihiu Beds (Reichenbacher 2000: 15). Unfortunately, the holotype is lost. According to the short description given by Weiler 1929) (Tab. 2). the ost significant characters of P. praecursor are the dorsal Iin with 9 long rays (instead of 78 in /'. alzi). the nuber of rays in the anal fin (1112. instead of 1417 in P. alzi). and the position of the pelvic fins, which are situated nearer to the anal fin than to the pectoral fins (instead of the contrary in /'. alzi). ll is worth entioning that according to Weiler (1929). the caudal l of /'. praecursor is rather deeply furcated because this can be hardly expected with in the genus Prolebias S \i \ \<.i. Moreover, our reexaination of /'. cl. prae cursor fro ihe Ceriihiu Beds of Ihe borehole Gundershei 13. deterined by Weiler 1963). revealed thai this specien has two dorsal fins, the first one being ade of short thin spines. Consequently. P. cf. pruecursor cannot belong to the genus Prolebias. but ay be a species of the genus Heilrichas Pi 11 its which is widely distributed in the Main/ Basin and Ihe Hanau Basin (Micklich Ott Keller et al. 2002). Prolebias rhenanus Gaudani 1981 is known fro the Lower Oligocene of Kleinkes (near Baden) in the southern Upper Rhinegraben. This species, whose characters are suarized in Tab. 2. clearly differs fro P. alzi because of its saller anal fin consisting of 1213 rays (instead of 1417 in P. inalzi). and its pelvic fins, which are saller and situated nearer to the anal fin than to the pectoral fins (instead of the contrary in P. alzi). 7. Otolith variability ot Prolebias alzi n. sp. We have investigated the angle of the excisura in populations of P. alzi otoliths fro the Upper Cerithiu Beds and the overlying Rüssingen Foration (forer Inflata Beds). The ex cisura angle tends to be narrower in older speciens (Fig. 10). This fits well with the fact that the excisura angle is ost nar row (114 in the otolith fro the Middle Cerithiu Beds of Dudenhofen (Fig. 9). Also the antirostru displays changes in shape and diensions fro the older to the younger popula tions (Reichenbacher 2000: Fig. 17). Following the actualistic nube ol otoliths Fig. 10. Distribution patterns of excisura angles of Prolebias inalzi n. sp. Iro the Upper Cerithiu Beds (dark coluns, n 16) and the overlying RUssingen Foralïon (light coluns, n 18). studies on the Recent Mediterranean species Aphanius iberus (Valenciennes in Cuvier & Valenciennes) (Reichenbacher \ Sicnknechl 2001 such changes otolith orpholog) a; point to a genetic divergence between Prolebias alzi popula tions of different stratigraphic ages. Acknowledgeents We are grateful lo W. On (Offenbach), K. Weiss (Fischbach), and M. Keller (Franklin!), who ade available their newly collected cyprodiintitor fishes and donated the ost iportant speciens to the Senckenberg Museu. We are grateful lo Dr. N. Micklich (Hessisches Landesuseu Darstadt) and Prof. Dr. G. Stringer (The University of Louisiana at Monroe) for Iheir critical review of the anuscript. We thank Dr. G. Plodowski and Dr. T. Jellinek (both Forschungsinstitut und Naturuseu Senckenberg. Frankfurt/Main) l'or the loan of aterial. Sincere thanks go to D. Serrette (Paris) for the pho tographs of the fish skeletons and to J. Du (Paris) lor preparing the illustra tions of Figs. 45. SEMpictures were ade at the Laboratoriu für Elektro nenikroskopie of the University Karlsruhe, we thank V. Zibat for his assislance Ihi studv 77.is kindly supported b) ilu Deutsche Forschungsgeein schaft (Re 1113/61). REFERENCES A(,\ssi/. L. 1839: Recherches sur les Poissons fossiles. 5(12) Neuchâlel (Pelilpierre). Bkst. G. 1975: Feinstraligraphie der HydrobienSchichten (Unteriozän. Main/er Becken) Main/er geowiss. Mitt Mainz. ERASMO. G. D' 1929: L'Ittiofauna fossile di Senigallia. Atti della R. Accade ia delle Scienze fisische e ateatiche di Napoli G vi dant. J. 1981: Un nouveau Cyprinodontidae (Poisson téléostéen) de l'oligocène inférieur de Kleinkes (Pays de Bade. Alleagne): Prolebias rhenius nov. sp. Sci. Geol. Bull. 34(1) G \i d\. J. 1989: Découverte d'une nouvelle espéce de poissons cyprinodon tifores (Prolebias delphiiicnsis nov. sp.) dans l'oligocène du bassin de MontbrunlesBains (Dròe). Géol. éditerr Marseille On Prolebias es cri Ac. vssiz) fro the OligoMiocene of the Upper Rhinegraben 519

13 Gaudant. J : Prolebias hungaricus nov. sp.: une nouvelle espèce de Pois sons Cyprinodontidae des diatoites iocènes de Szurdokpüspöki (Coté de Nograd. Hongrie). Magyar All. Földt. Int. Evi Jel Gaudant. J. 1998: L'ichthyofaune des eaux continentales iocènes de Serbie (Yougoslavie): une revision. N. Jb. Geol. Paläont. Abh. 207(1) Gai Dani. J. & RoviraSendrôS. J. 1998: Découverte de la plus ancienne es pèce connue du genre Aphanius Nardo (Poissons téléostéens) dans le Miocène inférieur évaporilique de la Catalogne. Batalleria 8, Gaudant, J. & Reichenbacher, B. 2002: Anatoie et affinités des Prolebias aff. weileri von Salis (Poissons téléostéens. Cyprinodontidae) du Miocène inférieur à oyen du Randecker Maar (Wurteberg. Alle agne). Stuttgarter Beitr. Naturkde. B Keller. T.. Reichenbacher. B.. Gaudant. J. 2002: Erstnachweis von AtherinidenSkcletten (Pisces. Teleostei. Atherinifores) il Otolithen in situ aus den Unteren HydrobienSchichten (WiesbadenForation) des Mainzer Beckens, it eine Beitrag zur Sedientologie und Paläookolo gic. Cour. Forsch.Inst. Senckenberg Malz. FI. 1978: Vergleichendorphologische Untersuchungen an aquitanen FischOtolithen aus de Untergrund von Frankfurt a Main. Sencken berg. lethaea 59(4/6) Martini. E. 1987: Nannoplankton aus den Oberen HydrobienSchichten des Hanauer Beckens (Miozän). Geol. Jb. Hessen Meyer. H. v. 1834: Vorkoen des Lebias eyeri. Agass.. eines fossilen Fi sches, i Thone von Frankfurt a.m. Mus. Senckenbergianu Micklich. N. 1996: Sexual diorphis in a lower Miocene oronid? Pal. Z. 70(3/4) Ott. W. 2001: Baustellen: Fenster in die Erdgeschichte. 2. Mitteilung: Baustelle "Westhafen". Frankfurt a Main. Ber. Offb. Ver. Naturkde : Baustellen: Fenster in die Erdgeschichte. 3. Mitteilung: Neubau der UBahnstalion Senckenberganlage. Frankfurt a MainBockenhei. Eine ergiebige Fundstelle fossiler Fischskelette in den Oberen Hydrobi enschichten. Ber. Offb. Ver. Naturkde 103 (in press). Parenti. L. R. 1981: A phylogenetic and biogeographic analysis of Cyprinodontifor fishes (Teleostei. Atherinoorpha). Bull. Aer. Mus. Nat. Hist. 168(4) Reichenbacher. B. 2000: Das brackischlakustrine Oligozän und Unter Miozän i Mainzer Becken und Hanauer Becken: Fischfaunen. Paläoökologie. Biostratigraphic Paläogeographie. Cour. Forsch.Inst. Senckenberg Reichenbacher. B. & Keller. T. 2002: Neudefinition von stratigraphischen Einheiten i Tertiär des Mainzer und Hanauer Beckens (Deutschland. Oligozän Miozän). Teil 2: WiesbadenForation [= Untere Hydrobien Schichten]. Mainzer geowiss. Mitt Reichenbacher. B. & MÖDDEN, C. 1996: Biostratigraphie und Paläoökologie aufgrund von FischOtolithen in den Oberen Cerithienschichten (Unter Miozän) bei Göllhei (Mainzer Becken). Mainzer geowiss. Mitt (1. Reichenbacher, B. & Sienknecht, U. 2001: Allopatric divergence and ge netic diversity of recent Aphanius iberus and fossil Prolebias eyeri (Teleostei. Cyprinodontidae) fro southwest and western Europe, as in dicated by otoliths. Geobios 34(1) Reichenbacher. B. & Weidann. M. 1992: FischOtolithen aus der oligo /iozänen Molasse der WestSchweiz und der HauteSavoie (Frankreich). Stuttgarter Beitr. Naturkde. B Sauvage. H.E. 1874: Notice sur les Poissons tertiaires de l'auvergne. Bull. Soc. Hist. nat. Toulouse 8 ( ) SCHÄFER, P. & Kadolksy. D. 2002: Neudefinition von stratigraphischen Ein heiten i Tertiär des Mainzer und Hanauer Beckens (Deutschland. Oligozän Miozän). Teil I: OberradForation Obere Cerithien schichten. oberer Teil) und RUssingenForation [= /n/7afaschichten]. Mainzer geowiss. Mitt Sei Hl. R.P. 1960: Osteology and Phylogeny of Oviparous Cyprinodont Fishes (Order Cyprinodontifores). Diss. Univ. Florida Sienknecht. U. & Reichenbacher. B. (in prep.): Otolith affinities of Valen cia Myers and Aphanolebias Reichenbacher & Gaudant and phyloge netic conclusions. ViLLWOCK W. 1999: Biogeography of the Cyprinodonlifor fishes (Teleostei: Cyprinodontidae) of the Mediterranean region. In: Generalität Valenciana. (Ed. by Conselleria de Medio Abiente) Peces Ciprinodóntidos Ibéricos Fartet y Saaruc. Monografia: Weiler. W. 1929: Prolebias praecursor sp. n. aus den Süsswasserschichten des Cyrenenergels i Mainzer Becken. Notizbl. Ver. Erdk. 5(12) Weiler. W. 1942: Die Otolithen des rheinischen und nordwestdeutschen Ter tiärs. Abh. Reichsat Bodenforsch. N. F Weiler. W. 1963: Die Fischfauna des Tertiärs i oberrheinischen Graben, des Mainzer Beckens, des unteren Maintales und der Wetterau. unter besonderer Berücksichtigung des Unteriozäns. Abh. Senckenberg. naturforsch. Ges Wiesner. E. 1967: Das Miozän i östlichen Mainzer Becken unter besonderer Berücksichtigung der Mikrofauna. Unpublished Thesis Geol Paläont. Inst.. Univ. Frankfurt/Main. Manuscript received February Revision accepted Septeber B. Reichenbacher & J. Gaudant