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Revision of the planktonic foraminifera Pseudothalmanninella klausi (LEHMANN, 1966) and Globotruncanella semsalensis (CORMINBOEUF DQGÀUVWVFDQQLQJ HOHFWURQPLFURVFRSH 6(0 LPDJHVRIWKHKRORW\SHV

Silvia SPEZZAFERRI1 & Michèle CARON1

Abstract The planktonic foraminifera Rotalipora klausi LEHMANN, 1966 and Globotruncana? semsalensis CORMINBOEUF, 1961, ZHUHÀJXUHG only by drawings and the specimens were never made publicly available, although deposited in the collections of the Department RI*HRVFLHQFHV)ULERXUJ6ZLW]HUODQG:HSUHVHQWKHUHIRUWKHÀUVWWLPHWKHLU6FDQQLQJ(OHFWURQ0LFURVFRSHGRFXPHQWDWLRQDQG comparison with the original drawings. We complete the morphological descriptions and, accordingly, we emend the generic attribution of both species. 6LQFHLWVÀUVWGHVFULSWLRQRotalipora klausi has acquired a great importance as phylogenetic and biostratigraphic marker for the Late Albian. Its generic attribution is made according to the recent revision of the polyphyletic group of Rotaliporids as Pseudothalmanninella klausi. Globotruncana? semsalensisZDVÀUVWGHVFULEHGLQDULFKSODQNWRQLFIRUDPLQLIHUDODVVHPEODJHRI8SSHU&DPSDQLDQ7KLVVSHFLHVZDV disregarded for long time because of some disturbing resemblances with the Praeglobotruncana lineage. The present study proves the validity of this species and attributes it to the genus Globotruncanella.

.H\ZRUGV: Planktonic foraminifera, holotypes, Cretaceous, genus revision

Résumé Révision des espèces de foraminifères planctoniques Pseudothalmanninella klausi (LEHMANN, 1966) and Globotruncanella semsalensis (CORMINBOEUF   HW SUHPLqUHV LPDJHV GHV KRORW\SHV DX PLFURVFRSH pOHFWURQLTXH j EDOD\DJH 6(0   Les espèces des foraminifères planctoniques Rotalipora klausi LEHMANN, 1966 et Globotruncana? semsalensis CORMINBOEUF, 1961, ont pWpÀJXUpHVVHXOHPHQWSDUGHVGHVVLQVHWOHVVSpFLPHQVQ·RQWMDPDLVpWpDFFHVVLEOHVDXSXEOLFELHQTXHGpSRVpVGDQVOHVFROOHFWLRQVGX Département de Géosciences, , Suisse. Nous présentons ici pour la première fois leur documentation au Microscope Electronique à Balayage accompagnée des dessins originaux. Nous complétons les descriptions morphologiques et, en accord avec ces observations, nous émendons les attributions génériques des deux espèces. Depuis sa première description Rotalipora klausi a acquis une grande importance comme marqueur phylogénétique et biostratigraphique SRXUO·$OELHQVXSpULHXU6RQDWWULEXWLRQJpQpULTXHHVWIDLWHHQDFFRUGDYHFODUpYLVLRQUpFHQWHGXJURXSHSRO\SK\OpWLTXHGHV5RWDOLSRULGV à savoir Pseudothalmanninella klausi. http://doc.rero.ch Globotruncana? semsalensis a été décrite dans un riche assemblage de foraminifères planctoniques du Campanien supérieur. Cette espèce a été négligée pendant longtemps à cause de certaines ressemblances troublantes avec la lignée des Praeglobotruncana/·pWXGH présentée ici prouve la validité de cette espèce et la rattache au genre Globotruncanella.

0RWVFOpV Foraminifères planctoniques, holotype, Crétacé, révision des genres.

I. INTRODUCTION holotypes of all the Cretaceous species. Consequently, some disregarded and/or neglegted holotypes have been Presently, the Mesozoic Planktonic Foraminifera Working sought and some of them rediscovered in ancient Group (of which the 2 authors are members), chaired by collections. B. HUBER of the Smithsonian Institution, is engaged in The holotypes of Rotalipora klausi LEHMANN, 1966 and the compilation of the on-line dictionary of Mesozoic Globotruncana? semsalensis CORMINBOEUF, 1961 planktonic foraminifera in the framework of the Chronos deposited in the collections of Cretaceous planktonic initiative (US-NSF Project). The dictionary includes the foraminifera of the Department of Geosciences of the Scanning Electron Microscope (SEM) images of the University of Fribourg were neglected for long time.

1 University of Fribourg, Department of Geosciences, Earth Sciences, Chemin du Musée 6, CH-1700 Fribourg, . E-mails: [email protected], [email protected]

1 However, the increasing stratigraphic and phylogenetic circular outline, chambers slowly increase in size as importance of the two species recently urged the added; a beaded keel is present in all chambers of the last availability of their SEM images. We present here their whorl, it is visible but smoother in the last chamber; ÀUVW 6(0 GRFXPHQWDWLRQ DQG FRPSDUH WKHP ZLWK WKH spiral sutures are curved and raised except between the 3 original drawings of KLAUS, 1959 (according designation last chambers where they are radial and depressed. On the by LEHMANN, 1966) and CORMINBOEUF, 1961). This umbilical side, sutures are radial and depressed. revision proves the validity of both species and clarify 3HULXPELOLFDOULGJHVDUHZHOOGHYHORSHGRQWKHIRXUÀUVW their generic attribution. chambers; the fourth periumbilical ridge raises exceptionally on the suture with the previous chamber. 7KHXPELOLFDODUHDLVQDUURZWKHWKUHHÀUVWVXSSOHPHQWDU\ II. SYSTEMATICS apertures are placed deeply in the umbilical depression; the supplementary aperture of the last chamber is well Pseudothalmanninella klausi (LEHMANN, 1966) visible in the umbilical area. 3O,ÀJVDFDF This species is here attributed to the genus Pseudothalmanninella, based on the newly emended +LVWRU\The species was erected by LEHMANN, 1966, in GHÀQLWLRQRIWKLVJHQXVJLYHQE\GONZALEZ DONOSO et his study on the coastal basin of Tarfaya, Morocco, where al. (2007) as it possesses the characters of he documents a rich Late Albian-early Cenomanian Pseudothalmanninella. In particular, it displays a spiro- planktonic foraminiferal fauna found in the Puerto FRQYH[SURÀOHZLWKORZZKRUOH[SDQVLRQUDWHHLJKWWR &DQVDGRZHOO+HLGHQWLÀHGPDQ\VSHFLPHQVWKDWFRXOG nine chambers are present in the last whorl; the keel is be clearly distinguished from the well known species present in all the chambers of the last whorl; the umbilicus Thalmanninella ticinensis (GANDOLFI) because of their is narrow and bordered by periumbilical ridges on the more numerous chambers and higher trochospire. ÀUVWIRXUFKDPEHUVWKHVXWXUHVDUHUDGLDODQGGHSUHVVHG However, he designated as holotype of this new species all supplementary apertures are present in umbilical (that he called Rotalipora klausi) a specimen documented position. in a publication of KLAUS SO,,ÀJVDF ZKHUH 5HPDUNVP.klausi is considered by GONZALEZ DONOSO LW ZDV LGHQWLÀHG DV Rotalipora (Thalmanninella) et al. (2007), as an intermediate form between P. ticinen- multiloculataDQGMXVWLÀHGKLVGHFLVLRQVRWKDW´«LWLV sis (GANDOLFI) and P. tehamaensis (MARIANOS & more judicious to choose a more accessible type-locality, ZINGULA). that is not the case for the Puerto Cansado well.” &RPSDULVRQThis species is larger, with more numer- The type specimen of KLAUS was deposited in the ous chambers and with a higher trochospire with respect collection of Cretaceous planktonic foraminifers at the to Pseudothalmanninella ticinensis. Although umbilical Department of Geosciences, Fribourg, and is documented sutures are depressed, P. klausi differs from P. ticinensis here. by having periumbilical ridges (not represented in the 7\SH UHIHUHQFH Rotalipora klausi LEHMANN 1966, drawing of KLAUS, but clearly visible in the SEM photo- http://doc.rero.ch designated in KLAUS,  SO ,, ÀJ DF VSHFLPHQ graphs of the holotype). It differs from P. tehamaensis in .F DQG ÀJXUHG XQGHU WKH QDPH ´Rotalipora having depressed umbilical sutures although periumbili- (Thalmanninella) multiloculata (MORROW)”. FDOULGJHVDUHSUHVHQWRQWKHIRXUÀUVWFKDPEHUVRIWKHODVW +\SRW\SHV Illustrated in LEHMANN,ÀJVDMN whorl. OPQDQGSOÀJ 7\SH ORFDOLW\ La Forclaz Pass, slope of the Dent de 5HSRVLWRU\ The holotype is now deposited at the Natural Bourgoz, Prealpes Medianes of Fribourg, Switzerland. History Museum of Basel with the reference number $JHLate Albian. NMB-C39016. The repository of the LEHMANN·V $FFRPSDQ\LQJ DVVHPEODJH Pseudothalmanninella hypotypes from Morocco is unknown. ticinensis, Thalmanninella appenninica, Planomalina 2ULJLQDO GHVFULSWLRQ Biconvex test formed by three EX[WRUÀand other species as shown in Fig. 1. trochospiral whorls, more convex on the spiral side than 5HIHUHQFHELR]RQHTh. appenninica Zone. on the umbilical side. Eight to nine chambers are present *HRJUDSKLF GLVWULEXWLRQ Alps (Western Tethys); in the last whorl and have roughly the same size. The test Morocco (North Atlantic) and Tunisia (Tethys). outline is moderately lobate. Umbilical sutures are radial with accessory aperture located in the narrow umbilicus. Globotruncanella semsalensis (CORMINBOEUF, 1961) The primary aperture is interiomarginal, umbilical- 3O,ÀJVDFDF3O,,ÀJVDFDFDF extraumbilical. See KLAUS·VGUDZLQJVRIWKHKRORW\SH 3O,ÀJVDF +LVWRU\ The samples studied by CORMINBOEUF (1961) *HQXVUHYLVLRQ The revision of the genus of this species ZHUHWDNHQLQWKH´:LOGÁ\FKVXEDOSLQµ=RQHRXWFURSSLQJ LVEDVHGRQREVHUYDWLRQRIWKHKRORW\SH 3O,ÀJVDF  in the Semsales region, , Switzerland. Nine chambers are present in the last whorl and are The age of this formation is Middle Oligocene in age. arranged in a high trochospire. The spiral side shows a However, the section studied by CORMINBOEUF (1961)

2 Planktonic Foraminiferal Polarity Stages Chron Praegl. stephani Age (Ma) R. cushmani Zones Praegl. delrioensis Th. balernaensis Th. appenninica Pseudoth. tehamaensis Th. globotruncanoides Th. reicheli Th. greenhornensis Pseudoth. ticinensis Pseudoth. klausi R. montsalvensis Th. praebalernaensis T. roberti T. buxtorfi P. Pseudoth. subticinensis

95 R. cushmani

Th. reicheli

Th. globotru- Cenomanian ncanoides

99.6 100 Th. appen- ninica

Th. ticinensis

Th. subticin. Albian

T. prae- 105

http://doc.rero.ch ticinensis Cretaceous Normal Polarity Super Chron (Cretaceous Quiet Zone)

Fig. 1: Distribution of genera and species of planktonic foraminifera: Pseudothalmanninella klausi (LEHMANN) from Late Albian, according to KLAUS (1959), GONZALEZ DONOSO et al. (2007), ROBASZYNSKI &CARON (1995) and PREMOLI SILVA & VERGA  . Time scale from OGG et al.   T. = Ticinella; P. = Planomalina; Pseudoth. = Pseudothalmanninella; Th. = Thalmanninella; R. = Rotalipora; Praegl. = Praeglobotruncana,

situated in an old state quarry 1.5 km north-east of green marls, just below the B. mucronata layer, contains ´5XLVVHDXGHOD&LHUQHµLVDQROLVWROLWKDERXWPWKLFN a rich association of Upper Campanian planktonic and comprises an alternation of white-grey micritic foraminifera from the G. aegyptiaca-G. falsostuarti Zone. limestones and calcareous marls (CHUNGKHAM & CARON, CORMINBOEUF (1961) described Globotruncana ? 1996, p. 503). In the lower part of the section limestones semsalensis from this level. are predominant and contain an assemblage including The original material studied by CORMINBOEUF (1961) Belemnitella mucronata (SCHLOTH) var. minor JELETSKY was deposited in the collection of Cretaceous planktonic and Belemnitella cf. langei JELETZKY, which corresponds foraminifers at the Department of Geosciences, Fribourg. to the Bostrychoceras polyplocum ammonite Zone, The collection was recently revised following the according to CORMINBOEUF (1961). The level of grey- suggestion of the Mesozoic Planktonic Foraminifera

3 Working Group with the aim to search and complete the apertures in the back. The infralaminal apertures (main data set for all type specimens present in the collection and secondary) may have – or not – a thin lip. Each and never made available by the author. tegillum partly or totally extends over the tegillum of the 7\SHUHIHUHQFH: Globotruncana? semsalensis CORMIN- preceding chamber, the tegillum of the last chamber BOEUF,3O,,ÀJVDFVDPSOH& sometimes seals the whole umbilicus. Secondary 3DUDW\SHV CORMINBOEUF, 3O,,ÀJVDFDF infralaminal apertures are not present. See drawings of VDPSOH& CORMINBOEUF  3O,ÀJVDF 5HSRVLWRU\ The holotype is presently deposited at the Genus revision: The revision of the genus of this species Natural History Museum of Basel with the reference LVEDVHGRQWKH6(0LPDJHVRIWKHKRORW\SH 3O,ÀJV number NMB- C39017, together with the two paratypes DF DQGRQGUDZLQJVRIWKHSDUDW\SHV 3O,,ÀJVDF illustrated by CORMINBOEUF   LQ 3O ,, ÀJV DF 3a-c) and 3a-c, and with a washed residue from a sample taken Six chambers are present in the last whorl. Chamber in the same type level and containing a rich assemblage arrangement is trochospiral with very variable height (Pl. of planktonic foraminifera. ,,ÀJV ,WLVORZLQWKHKRORW\SHSURÀOHWRKLJKLQWKH 2ULJLQDOGHVFULSWLRQ VLPSOLÀHG The test is trochoidal second paratype. The spiral side shows a lobate outline; and lobate. Generally six chambers are present in the last chambers are slowly increasing in size as added; strong whorl. They are compressed and slowly increasing in size SXVWXOHVDUHSUHVHQWRQWKHVXUIDFHRIWKHÀUVWFKDPEHUV as added. The last-formed chamber can be larger, the of the last whorl, thicker and concentrated along the same size or even smaller than the penultimate chamber. peripheral row; this ornamentation disappears Pustules are rounded, slightly elongate, or spine-like, progressively on the last two chambers, the last one being they are densely present on the two sides of the test. On always smooth, but exceptionally smaller as in the WKH ÀUVW FKDPEHUV RI WKH ODVW ZKRUO WKH\ DUH WKLFN KRORW\SH 3O,,ÀJVDF  gradually becoming thinner in the following chambers so On the two inner whorls, the peripheral double row of that the last chamber(s) may be more or less smooth. The pustules underlines the raised spiral sutures, continuing spiral side is convex. In the second whorl, chambers have RQ WKH  ÀUVW FKDPEHUV RI WKH ODVW ZKRUO WKH  ODVW FKDPEHUVEHLQJLQÁDWHGWKHVSLUDOVXWXUHVDUHGHSUHVVHG DÁDWRUVOLJKWO\FRQYH[VXUIDFHZLWKWKLFNURXQGHGVSLN\ On the umbilical side, sutures are radial and depressed. or slightly elongate pustules concentrating along the 7KHSXVWXOHVDUHPRUHGHQVHO\GLVWULEXWHGRQWKHÀUVW periphery and roughly shaping spiral and septal ridges. chambers of the last whorl, the last one being smooth. This ornamentation often remains present in the two or The umbilicus is one third of the test diameter; smaller WKUHHÀUVWFKDPEHUVRIWKHODVWZKRUOWKHQLWGLVDSSHDUV DQGGHHSHULQWKHKLJKO\VSLURFRQYH[SDUDW\SH 3O,,ÀJ resulting in septal sutures depressed, radial, or curved in 3c). The umbilical area is covered by a complex system frontward direction. The spiral side of the chambers of of portici; above the primary aperture, each chamber this last whorl is weakly to moderately convex. On the develops a large porticus towards the umbilicus, extending XPELOLFDOVLGHFKDPEHUVDUHVWURQJO\LQÁDWHGVXWXUHVDUH over the porticus of the preceding chamber. Two or three radial and depressed. Strong and thick pustules are present successive portici are distinct in the umbilical area, http://doc.rero.ch RQWKHÀUVWFKDPEHUVRIWKHODVWZKRUOHLWKHUUDQGRPO\ particularly well developed in the paratype shown in Pl. distributed or aligned in the coiling direction or ,,ÀJFLQZKLFKWKH\UHVHPEOHDWHJLOOD perpendicular to that direction and, in this case, appearing Remarks: According to the characteristic features well like short costae spreading out from the umbilical edge visible on the holotype and paratype specimens, this toward the margin. In some specimens, one or two rows species is now attributed to the genus Globotruncanella. of pustules set in the coiling direction, become stronger The thick rounded pustules concentrating along the RQ WKH WZR RU WKUHH ÀUVW FKDPEHUV RI WKH ODVW FRLO DQG periphery and roughly shaping spiral and septal ridges double or triple the already existing peripheral row. The and the presence of large portici covering the umbilicus primary aperture is umbilical but it often extends slightly (and not of a tegilla), places this species as an intermediate toward the edge. It is covered by a tegillum directed between Globotruncanella havanensis (Voorwijk), with toward the umbilicus, each tegillum is provided with a less pustulous wall texture and G. pshadae (KELLER) main aperture in the front and with one or two secondary with keeled periphery.

Plate I

Figs 1a-c: Pseudothalmanninella klausi (LEHMANN), SEM images of the holotype. Figs 2a-c: KLAUS·V  GUDZLQJRIWKHKRORW\SH Figs 3a-c: Globotruncanella semsalensis (CORMINBOEUF  6(0 LPDJHV RI WKH KRORW\SH )LJV DF CORMINBOEUF·V (1961) drawing of the holotype. a = Spiral view, b = side view, c = umbilical view.

4 http://doc.rero.ch

5 &RPSDULVRQGlobotruncanella semsalensis, holotype VKRZLQJODUJHSRUWLFL ÀUVWO\GHVFULEHGDV´WHJLOODµE\WKH and two paratypes (Pl. II), differ from the members of the author himself, CORMINBOEUF, 1961). Praeglobotruncana delrioensis - P. stephani - P. gibba 7\SHORFDOLW\´&DUULqUHGHO¶(WDWµ6HPVDOHV&DQWRQRI lineage in having strong pustulosous chamber surfaces, Fribourg, Switzerland. sometimes forming a meridional pattern, in having the $JHLate Campanian. periphery which never develops a double keel and in $FFRPSDQ\LQJ DVVHPEODJH Rugoglobigerina ssp.,

Planktonic Foraminiferal Age (Ma) Stages Polarity Chrons Zones Gta. stuarti A. intermedius A. mayaroensis G. aegyptiaca R. macrocephala G. linneiana G. bulloides G. falsostuarti Gta. stuartiformis Rad. calcarata Glla. pshadae Glla. semsalensis R. rugosa R. hexacamerata Gan. gansseri Glla. havanensis C. fornicata 65 65.5 C29 Abathomphalus C30 mayaroensis

Maastr. C. contusa / C31 R. fructicosa 70 70.6 G. gansseri C32

G. aegyptiaca

75 G. havanensis G. calcarata

G. ventricosa

Campanian C33 http://doc.rero.ch

80

G. elevata

83.5

Fig. 2: Distribution of genera and species of planktonic foraminifera: Globotruncanella semsalensis (CORMINBOEUF) from Upper Campanian, according to CORMINBOEUF (1961), ROBASZYNSKI & CARON (1995) and PREMOLI SILVA & VERGA  . Time scale from OGG et al.  Glla. = Globotruncanella; R. = Rugoglobigerina; G. = Globotruncana; Gta. = Globotruncanita; Rad. = Radotruncana; Gan. = Gansserina and A. = Abathomphalus.

Plate II

Figs 1a-c: Globotruncanella semsalensis (CORMINBOEUF, 1961), drawings of the holotype. Figs 2a-c: Globotruncanella semsalensis (CORMINBOEUF, 1961), drawings of the 1st paratype. Figs 3a-c: Globotruncanella semsalensis (CORMINBOEUF, 1961), drawings of the 2nd paratype.

6 http://doc.rero.ch

7 Globotruncana linneiana, G. bulloides, G. falsostuarti, REFERENCES G. aegyptiaca, Globotruncanita stuartiformis and other species as shown in Fig. 2. CORMINBOEUF, P. (1961) - Association de Belemnitella et de 5HIHUHQFH ELR]RQH G. aegyptiaca/G. falsostuarti Globotruncanidae dans le Campanien supérieur des Zones. Alpettes (Préalpes externes fribourgeoises). Eclogae *HRJUDSKLFGLVWULEXWLRQ Alps (Western Tethys). Geologicae Helvetiae   CHUNGKHAM, P. & M. CARON (1996) - Comparative study of a late Maastrichtian foraminiferal assemblage, Semsales :LOGÁ\VFK=RQH6ZLW]HUODQGDQG8NKUXO0HODQJH=RQH III. CONCLUSIONS India. Revue de Paléobiologie*HQqYH   GONZALEZ-DONOSO, J.-M., D. LINARES, M. CARON & F. :HGRFXPHQWWKHÀUVW6(0LPDJHVRIWKHKRORW\SHVRI ROBAZYNSKI (2007) - The rotaliporids, a polyphyletic Rotalipora klausi LEHMANN, 1966 and Globotruncana? group of Albian-Cenomanian planktonic foraminifera. semsalensis CORMINBOEUF, 1961, and we compare them Emendation of Genera. Journal of Foraminiferal Research, with the original drawings. Based on these new Washington, 37(2): 175-186. observations we are able to revise the two genera and KLAUS, J. (1959) - Le «Complexe schisteux intermédiaire» dans le synclinal de la Gruyère (Préalpes médianes). attribute them to Pseudothalmanninella and Eclogae Geologicae Helvetiae, 52(2): 753-851. Globotruncanella respectively. The revision has been LEHMANN, R. (1966) - Les Foraminifères pélagiques du Crétacé QHFHVVDU\ EHFDXVH VLQFH WKHLU ÀUVW GHVFULSWLRQ ERWK du bassin côtier de Tarfaya. 1. Planomalinidae et Pseudothalmanninella klausi and Globotruncanella Globotruncanidae du sondage de Puerto Cansado (Albien semsalensis have acquired a great importance as supérieur, Cénomanien inférieur). Notes et Mémoires du phylogenetic and biostratigraphic markers, for the Late Service Géologique du Maroc, 175: 153-167. Albian (P. klausi) and for the Late Campanian (G. OGG, J., F.P. AGTERBERG &F.M.GRADSTEIN (   7KH semsalensis). Cretaceous Period. In : GRADSTEIN, F.M., J. OGG & A. SMITH (Eds). A geologic time scale, Cambridge University 3UHVV PREMOLI SILVA, I. & D. VERGA  3UDFWLFDO0DQXDORI ACKNOWLEDGMENTS Cretaceous Planktonic Foraminifera. In:VERGA, D. & R. RETTORI (Eds). International school on Planktonic We warmly thank all the members of the Mesozoic Foraminifera. Universities of Perugia and Milano, Planktonic Foraminifera Working Group for the fruitful 7LSRJUDÀD3RQWHIHOFLQR3HUXJLD ,WDO\ S GLVFXVVLRQ DQG HQFRXUDJHPHQW WR GHVFULEH ´QHJOHFWHG´ ROBASZYNSKI F. & M. CARON (1995) - Foraminifères holotypes. Thanks to C. NEURURER (Fribourg) for planctoniques du Crétacé: Commentaire de la zonation assistance in SEM operation. Europe-Méditerranée. Bulletin de la Société Geologique de , 166(6): 681-692. http://doc.rero.ch

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