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Species Planorotalites pseudoscitula Glaessner 1937



Diagnosis / Definition:
Pearson et al. (2006):
DESCRTPTION. Type of wall: Normal perforate, nonspinose, weakly muricate. Test morphology: Peripheral outline oval to subcircular, weakly lobulate, coiled in low trochospire, weakly biconvex; in umbilical view 6-8 subtriangular chambers, compressedlflattened along peripheral margin, intercameral sutures radial (in early part of last whorl) to slightly curved in younger chambers, depressed in terminal 2-3 chambers, umbilicus narrow, shallow, aperture a low, umbilical-extraumbilical arch bearing a distinct lip; in spiral view 15- 18 trapezoidal chambers arranged in 2 1/2-3 whorls, sutures essentially flush with test chambers, distinctly curved, early part of test strongly muricate, elevated; in edge view test is biconvex, with distinct imperforate keel which extends at least to ante- or preantepenultimate chamber, test surface penetrated by normal-sized pores which do not open into distinct pore pits, aperture a typical umbilical-extraumbilical, low slit with distinct lip which extends to the periphery. Size: Typical dimension given by Glaessner (1937, p. 32): 0.2-0.25 mm.
Discussion / Comments:
Toumarkine & Luterbacher (1985):
This small species (0.2 to 0.3 mm) is lenticular, with 5 to 7 tightly coiled chambers in the last whorl. The periphery is circular, slightly lobate and with a faint keel. The surface of the test is rather smooth, but the surface of the chambers of the initial part of the last whorl may be somewhat rugose. The perforation is coarse in the older part of the test, fine in the youngest chambers.
Pearson et al. (2006):
DISTINGUISHING FEATURES.- The diagnostic features of Planorotalites pseudoscitula are its small, weakly biconvex and muricate, distinctly carinate test (see also under P capdevilensis). DISCUSSION- Berggren (1968, 1977), McGowran (1968), Schmidt and Raju (1973), Hillebrandt (1976) and Blow (1979) placed renzi Bolli in the synonymy of pseudoscitula Glaessner and in this most workers have followed them for the past 20 years. The supposed synonymy of these two forms as well as G. pseudoscitula elongata was suggested to one of us (WAB) by Gohrbandt as early as the mid-1960s based on examination and illustrations of a topotype of G. pseudoscitula elongata from Il 'skaya, northern Caucasus (sent to him by Glaessner) and 3 topotype specimens of renzi from Trinidad. Pertinent observations made at the time by Gohrbandt (pers. comm.) include the following: 1. "G. elongata has a more conical last chamber in axial profile (than G. renzi). G. renzi has a rather more acute periphery on the later chambers. However, G. pseudoscitula elongata shows signs of a 'keel'; and the difference is a minor one. There are no other significant differences, so far as can be seen, in the surface texture of the two forms. The external appearance suggests that oriented thin sections would show the same wall structure". 2. In spiral view the specimen of G. pseudoscitula elongata resembles the type figure of G. pseudoscitula s.s., whereas the axial profile resembles the variety elongata. 3. Russian workers no longer recognized elongata as a formal variety. In gross form G. renzi resembles the type figure of G. pseudoscitula very closely indeed. The similarity in surface texture has been confirmed; it had been strongly suspected from published data. The SEM was unavailable to Gohrbandt at the time he made his observations with the light microscope. SEM illustrations have shown that there is a difference in wall texture in the two forms, that of renzi (=capdevilensis, see above) being considerably more coarsely muricate. Therefore, we have refrained from concluding that G. renzi/ capdevilensis is a junior subjective synonym of G. pseudoscitula (see above). 4. Gohrbandt also observed that G. pseudoscitula (which he regarded as conspecific with G. renzi) had its lowest occurrence in the lower Eocene Globorotalia aragonensis Zone (distinctly lower than the base of Zone P10 as given by Bolli, 1957b; cf. Berggren, 1960 who had noted earlier an extension into lower Eocene in Nigeria as well). However, Blow (1 979) presciently observed that earlier (early Eocene) and later (middle Eocene) forms of this plexus are characterized by a coarsening of the muricate wall texture and gradual enlargement of pores (similar observations were made on this plexus by Benjamini, 1980). Blow (1979) was also correct in surmising that the type level of G. pseudoscitula Glaessner (and of G. elongata Glaessner) is of early (not middle) Eocene age as most investigators had thought. Indeed, the type level is from what was subsequently designated (Subbotina, 1953) the "Zone of compressed globorotaliids" (the Abazin Fm.), stratigraphically equivalent to Zones E4, ?E5. Both he (Blow, 1979, p. 898) and Bolli (1957b, p. 168) gave the stratigraphic range of G. pseudoscitula s.s. (= G. renzi) as Zones E81 E13 equivalents. However, in his text Blow (1979) referred to stratigraphically earlier morphotypes as Globorotalia cf. pseudoscitula with a range of Zones P7-P9; ? P10 (=E5-E7; ?E8) equivalents. Globorotalia renzi and G. capdevilensis partim were regarded as junior synonyms of G. pseudoscitula s.l.. Blow (1979) suggested that the wall texture pattern changes were one of degree rather than kind and that Globorotalia pseudoscitula s.s. (rather than Globorotalia cf. pseudoscitula based on his earlier, mistaken belief that the type level of G. pseudoscitula was middle Eocene) might be appropriate for the older (early Eocene) forms and G. renzi for the younger (middle Eocene) forms, while retaining the designationpseudoscitula s.l. for the entire plexus. Accordingly it is clear that Gohrbandt had made his observations on elongata (from the basal Eocene Zone E3-ES), and on renzi from upper middle Eocene Zone E 1 2-E 13. At the same time Blow (1 979) considered that Globorotalia elongata Glaessner is essentially unrelated to the G. pseudoscitula/renzi plexus. This was based upon the following lines of evidence: 1. Blow (1979) had submitted several specimens (subsequently illustrated as figures 6 and 7 of his plate 116 from [his] Zone P7 [=E51 of the Kilwa area, Tanzania), to Martin Glaessner who confirmed their identification as Globorotalia elongata; this led Blow (1 979, p. 890) to assert that (Paleocene) records of G. elongata by Loeblich and Tappan (1 957) and Bolli (1 957a) did not correspond to Glaessner's elongata. 2. G. elongata Glaessner is characterized by an acute to subacute peripheral margin and "nearly equally biconvex test". Furthermore, elongata is a minute (0.15-0.2 mm; Glaessner, 1937; 0.167-0.249 mm: Blow, 1979) form. Confusion with G. pseudomenardii was not possible according to Blow because of the less ventrally inflated chamber surfaces and smaller umbilicus in pseudomenardii, "which, especially in stratigraphically younger specimens are virtually closed (Blow, 1979, p. 891). 3. G. elongata was said to differ from G. pseudoscitula in having a "more open and lax trochospire so that the later chambers increase in size more rapidly than in pseudoscitula" (Blow, 1979, p. 891). In the course of our work we have been unable to recognize a clear differentiation between morphotypes of elongata and pseudoscitula, considering the minor differences in coiling to be of degree rather than kind, and, accordingly, we include elongata in the synonomy of pseudoscitula. Phylogenetic relationships within this group remain enigmatic. McGowran (1968, lineage 6) and Berggren (1968, lineage 2), in trying to lay a phylogenetic foundation for distinguishing different clades of Paleogene "globorotaliids", recognized a morphologic and stratigraphic separation between Globanomalina and Planorotalites, although the former reserved Globanomalina for the planispiral forms (now referred to Pseudohastigerina) in the erroneous belief that the type species of Globanomalina (G. ovalis Haque) is planispiral (cf. Berggren and others, 1968; Banner, 1989, p. 171, 173). Owing to the superficial similarity among smooth-walled, finely perforate forms in the Paleocene the generic names Globanomalina and Planorotalites have been used interchangeably for Paleocene and lower Eocene taxa. Globanomalina has been adopted by the Working Group for the smoothwalled Paleocene group typified by G. ovalis and having its origin in G. archeocompressa in the basal Danian (Olsson and others, 1999; see also Berggren and Norris, 1997; Fleisher, 1974, p. 1017). Schmidt and Raju (1973) presented a detailed analysis of the morphology and evolution of the pseudoscitula plexus. Pertinent observations/conclusions include the following: 1. Globorotalia planoconica Subbotina is regarded as the stem form of the lineage and characterized by a small, finely perforate, unkeeled planoconvex test. 2. G. pseudoscitula Glaessner, 1937 is interpreted as a small, compressed, distinctly (coarsely) perforate, biconvex form with subcircular equatorial periphery and faintly keeled margin. G. pseudoscitula elongata Glaessner, 1937 is interepreted to represent varietal forms that develop a large, lobate last chamber. Globorotalia renzi is considered a junior synonym of G. pseudoscitula. 3. G. palmerae is considered to have evolved from pseudoscitula by "prolongation of the chambers at the axial periphery to a radially elongate shape terminating in a spine"(Schmidt and Raju, 1973, p. 179). 4. Globorotalia planoconica is shown to have originated in the G. velascoensis Zone (Zone P5 of this paper); G. pseudoscitula in the G. subbotinae Zone (Zone E3-E4 of this paper) and G. palmerae in the eponymous zone (i.e., within Zone E7). It would seem that these subtle differences in wall texture are at the root of the disparate lower range given for the pseudoscitula/renzi group, which spans the interval of Zones P5-E6 (Hillebrandt, 1962; Samuel, 1972; Benjamini, 1980; Fleisher, 1984; Toumarkine and Luterbacher, 1985). Benjamini (1980) suggested that the transition from the relatively larger, typical pseudoscitula with depressed sutures to the relatively small, coarsely perforate, flush sutured typical renzi took place in the mid-part of the G. palmerae Zone (= Zone E7 of this work) in the Avedat Group of the northern Negev, Israel. Subbotina (1953) had earlier noted that pseudoscitula extends down into the highest levels of the "Zone of rotaliid-like globorotaliids" (=Zones P213) in the Elburgan Formation of the North Caucasus, but we believe that these references are to members of the superficially similar Igorina pusilla group. In like manner, Loeblich and Tappan (1957) identified forms as pseudoscitula in the mid-late Paleocene of the Gulf Coast that, while superficially similar to the Eocene morphotypes, belong to the distinctly different group of the albearillaevigata plexus (Berggren, 1968). If the finely perforate, superficially smoothwalled (but wealtly muricate) to weakly cancellate, keeled, compressed, test of latest Paleocene-early Eocene pseudoscitula morphotypes are considered ancestral to the late early-middle Eocene, biconvex, keeled, coarsely perforate, muricate renzi, derivation from a juvenile morozovellid such as Morozovella occlusa would appear logical (cf. Berggren 1968; McGowran, 1968; Hillebrandt, 1 976 for earlier, alternate interpretations). In the course of our studies we have found that pseudoscitula has its lowest/earliest occurrence at the base of Zone P5 (i.e, just above the H0 of Globanomalina pseudomenardii). The specimen illustrated by Shutskaya (1956, pl. 4, figs. 5a-c) from the Cherkessk Fm. of the northwest Caucasus is excluded from this taxon. It would appear to be a relatively large (maximum diameter 0.43 mm) robust form that was compared by Shutskaya with Globorotalia subbotinae, G. nartanensis and G. spinulosa and was recorded from the G. subbotinae Zone (=Zone E3-4 of this paper), whereas P pseudoscitula is a relatively small form (with a maximum diameter of about half that of Shutskaya's specimen; Bolli's [1957a] holotype of Globorotalia renzi has a maximum diameter of 0.23 mm) and was shown to have its lowest occurrence considerably higher, in Zone P 10. The broadly rounded, seemingly densely muricate form illustrated by Hillebrandt (1962) from Zone P6 (equivalent) appears not to be referable to pseudoscitula/renzi as identified here and is more likely an acarininid referable to esnehensis (or a related form). PHYLOGENETIC RELATIONSHIPS- Unresolved; probably evolved from a juvenile morozovellid (? M. occlusa) by neoteny. STRATIGRAPHIC RANGE.- Zone P5-E7. GEOGRAPHIC DISTRIBUTION.- Ranges from tropics to temperate regions; central equatorial Pacific (ODP Site 865), New Jersey Coastal Plain (Bass River Borehole where it is a common form in Zone El), Tethyan deposits of northern Africa (Egypt where it appears in the lower part of the Esna Shale Fm. in basal Zone P5) and northern Caucasus; not reliably reported (to date) from high austral or northern latitudes to our knowledge. STABLE ISOTOPE PALEOBIOL0GY.- No data available.
Systematics:

15
 Classis Foraminifera
  Genus Planorotalites
   Species Hantkenina alabamensis
    Species Planorotalites pseudoscitula

35
  Ordo Foraminiferida
   Superfamilia Globigerinaceae
    Familia Truncorotaloididae
     Genus Planorotalites
      Species Planorotalites pseudoscitula
Synonym list:
Toumarkine & Luterbacher (1985):
1937 Globorotalia pseudoscitula Glaessner. - Glaessner : p.168 pl 38 figs 3a-c (type reference)
1957 Globorotalia renzi Bolli. - Bolli : p.168 pl 38 figs 3a-c
1985 Planorotalites pseudoscitula Glaessner. - Toumarkine & Luterbacher : p.118 figs 20.2-10
Pearson et al. (2006):
1937 Globorotalia pseudoscitula Glaessner. - Glaessner : p.32 text-fig. 3a-c [lower Eocene, holotype VNIGRI No. 4076, Lower Foraminiferal Beds, Suite F 1, Il'skaya, Northwest Caucasus]
1937 Globorotalia pseudoscitula var. elongata Glaessner. - Glaessner : p. 33 (German) p. 49 (Russian); text-fig. 3d-3f (p. 32) [Lower Foraminiferal Beds, Il'skaya, northwest Caucasus]
1947 Globorotalia pseudoscitula Glaessner. - Subbotina : 121-122 pl. 9; fig. 18-22 [Lower Eocene Globorotalia ex. gr. canariensis Zone, Abazin Fm., Khieu River section, northwest Caucasus]
1953 Globorotalia pseudoscitula Glaessner. - Subbotina : p.208 pl. 16; fig. 17a-c (reillustration of holotype No. 4076 of Glaessner, 1937) [Zone of compressed globorotaliids]; pl. 16, fig. 18a-c [Zone of thin-walled pelagic foraminifera, Foraminiferal Beds, Suite F2, North Caucasus]; pl. 17, fig. la-c [lower Eocene Zone of conical globorotaliids, Foraminiferal Beds, Suite F2, Khieu River section, North Caucasus]
? 1956 Globorotalia pseudoscitula Glaessner. - Shutskaya : p. 95, 96 pl. 4; fig. 5a-c [lower Eocene Globorotalia subbotinae Zone, Cherkessk Fm., near town of Nal'chik, northern Caucasus, FSU]
non 1957 Globorotalia pseudoscitula Glaessner. - Loeblich & Tappan : p. 193-194 pl. 46, fig. 4a-c; pl. 48, fig. 3ac; pl. 53, fig. 5a-c; pl. 59, fig. 2a-c; pl. 63, fig. 6a-c [=Igorina albeari (Cushman and Bermudez)]
non 1962 Globorotalia? (Globorotalia) pseudoscitula Glaessner. - Hillebrandt : p.129 pl. 12; fig. 12a-c [= ?Acarinina esnehensis (Nakkady)]
1972 Globorotalia pseudoscitula Glaessner. - Samuel : p. 193-194 pl. 51; fig. 2a-4c [upper Lutetian, Bakony Mountains, Hungary]
1973 Globorotalia pseudoscitula Glaessner. - Schmidt & Raju : p. 181, 182 pl. 1; fig. 3a-c, 4 [lower Eocene Globorotalia palmerae Zone, MNG-1, CC 18, Cauvery Basin, southeastern India]
1976 Planorotalites pseudoscitula Glaessner. - Hillebrandt : p.345 pl. 4; fig. 14 [lower Eocene Globorotalia palmerae Zone, Agost, Spain]
1977 Planorotalites pseudoscitula Glaessner. - Berggren : p.244 chart no. 3 (reillustrations from literature)
1979 Globorotalia (Globorotalia) pseudoscitula sensu lato Glaessner. - Blow : p.897 pl. 116, fig. 8-10 [lower Eocene Zone P7, Kilwa area, Tanzania, East Africa, referred to as G.(G.) cf. pseudoscitula]; pl. 173, fig. 1-8 [middle Eocene Zone P1 l, sample RS24, Kilwa area, Tanzania, East Africa]
1979 Globorotalia (Globorotalia) elongata Glaessner. - Blow : p.889 pl. 105, fig. 1-2, 4-6 [lower Eocene Zone P6, DSDP Hole 20C, South Atlantic Ocean]; pl. 116, fig. 6-7 [lower Eocene Zone P7, Sample RS 80, Kilwa area, Tanzania, East Africa]
1980 Planorotalites pseudoscitula Glaessner. - Benjamini : p.347 pl. 6; fig. 3-10 [Eocene Avedat Group, northern Negev, Israel]
p 1985 Planorotalites pseudoscitula Glaessner. - Toumarkine & Luterbacher : p.118 fig. 20.5 (holotype of Glaessner refigured); fig. 20.6 [lower Eocene Acarinina pentacamerata Zone, Richmond Fm., Jamaica]; fig. 20. 7-10 [Acarinina pentacamerata Zone, Cauvery Basin, South India, sample]
1988 Planorotalites pseudoscitula Glaessner. - Loeblich & Tappan : 477 pl. 518; fig. 6-8 [reillustration of holotype of Glaessner 1937]
1993 Planorotalites pseudoscitula Glaessner. - Lu & Keller : p.114 pl. 5; fig. 5-7 [lower Eocene Pseudohastigerina wilcoxensis (AP6a) Zone, ODP Hole 738C, Kerguelen Plateau, southern Indian Ocean]
1995 Planorotalites pseudoscitula Glaessner. - Lu & Keller : p.100 pl. 6; fig. 15-17 [lower Eocene Zone P6b, DSDP Site 577, Shatksy Rise, northwest Pacific Ocean]
2006 Planorotalites pseudoscitula Glaessner. - Pearson et al. : p.389 pl. 12.5; fig. 1-16
Was used in synonym list of:
Planorotalites capdevilensis Cushman & Bermudez 1949
Specimen:
VNIGRI collections, St. Petersburg, Inventory number: 4076
References:

Glaessner,M.F. (1937):
Planktonische Foraminiferen aus der Kreide und dem Eozän und ihre stratigraphische Bedeutung.
In: Studies in Micropaleontology Vol. 1(1) p. 27-46

Subbotina,N.N. (1947):
Foraminifery datskikh i paleogenovykh otlozhenii severnogo Kavkaza.
In: Mikrofauna neftyanykh mestorozhdenii Kavkaza, Emby I Srednei Azii Vol. 1 p. 39-160

Subbotina,N.N. (1953):
Iskopaemye foraminifery SSSR (Globigerinidy, Khantkenininidy i Globorotaliidy) . Trudy Vsesoyznogo Nauchno-Issledovatel'skogo Geologo-razvedochnogo Instituta (VNIGRI) Vol. 76 p. 296

Shutskaya,E.K. (1956):
Stratigrafiya nizhnikh gorizontov paleogena Tsentral'nogo Predkavkaz'ya po foraminiferam . Trudy Instituta Geologii Nauk Akademiya SSSR Vol. 164(70) p. 3-114

Loeblich,A.R. and Tappan,H. (1957):
Planktonic Foraminifera of Paleocene and Early Eocene Age from the Gulf and Atlantic Coastral Plains.
In: Studies in Foraminifera, United States National Museum Bulletin Vol. 215 p. 173-198

Bolli,H.M. (1957):
Planktonic Foraminifera from the Eocene Navet and San Fernando formations of Trinidad, B.W.I. . Bull. U.S. natl. Mus. Vol. 215 p. 155-172

Hillebrandt,v.A.. (1962):
Das Paleozän und seine Foraminiferenfauna im Becken von Reichenhall und Salzburg . Bayerische Akademie Wissenschaft, Mathematischen-Naturwissenschaften Klasse Vol. 108 p. 188 p.

Samuel,O.. (1972):
New species of planktonic foraminifers from the Paleogene of the west Carpathian in Slovakia (Czechoslovakia) . Zbornik Geologick"ck Vied Západné Karpaty Vol. 17 p. 165-221

Schmidt,R. and Raju,D.S.N. (1973):
Globorotalia palmerae Cushman & Bermudez and closely related species from the Lower Eocene, Cauvery Basin, South India . Proc. Koninkl. Nederlandse Akad. wetenschapen ser B Vol. 76 p. 167-178

Hillebrandt,A. (1976):
Los formainiferos planctonicos, nummulitidos y coccolitoforidos de las zona de Globorotalia palmerae del Cuisiense (Eoceno inferior) en el ES de Espana (Provincias de Murcia y Alicante) . Revista Espanol de Micropaleontologia Vol. 8 p. 323-394

Berggren,W.A. (1977):
Atlas of Paleogene planktonic foraminifera. Some species of the genera Subbotina, Planorotalites, Morozovella, Acarinina and Truncorotaloides.
In: Oceanic Micropaleontology Eds: Ramsay, A.T.S. p. 205-299

Blow,W.H. (1979):
The Cainozoic Globigerinida. 3 Vols p. 1413 pp

Benjamini,C.. (1980):
Plankton foraminiferal biostratigraphy of the Avedat Group (Eocene in the northern Negev, Israel) . Journal of Paleontology Vol. 54 p. 325-358

Toumarkine,M. and Luterbacher,H.P. (1985):
Paleocene and Eocene Planktic Foraminifera.
In: Plankton Stratigraphy p. 87-154

Loeblich,A.R. and Tappan,H. (1988):
Foraminiferal Genera and their Classification. 2 volumes. p. 970 pp

Lu,G. and Keller,G. (1993):
The Paleocene-Eocene Transition in the Antarctic Indian Ocean: Inference from Planktic Foraminifera . Marine Micropaleontology Vol. 21 p. 101-142

Lu,G. and Keller,G. (1995):
Planktic Foraminiferal Faunal Turnovers in the Subtropical Pacific during the Late Paleocene to Early Eocene . Journal of Foraminiferal Research Vol. 25 p. 97-116

Pearson,P.N.; Olsson,R.K.; Hemleben,C.; Huber,B.T. and Berggren,W.A. (2006):
Atlas of Eocene Planktonic Foraminifera. p. 1-513

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