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Chapter 14 TAXONOMY, BIOSTRATIGRAPHY, AND PHYLOGENY OF OLIGOCENE GLOBANOMALINIDAE (PSEUDOHASTIGERINA AND TURBOROTALIA) Paul N. Pearson 1 , Richard K. Olsson 2 , Silvia Spezzaferri 3 , and R. Mark Leckie 4 1 School of Earth and Ocean Sciences, Cardiff University, Main Building, Park Place, Cardiff, CF10 3AT, U.K. Email: pearsonp@cardiff.ac.uk 2 Department of Earth and Planetary Sciences, Busch Campus, Rutgers University, Piscataway, NJ 08854, U.S.A. Email: [email protected] 3 Department of Geosciences, Earth Sciences, University of Fribourg, Ch du Musée 6, 1700 Fribourg, Switzerland. Email: [email protected] 4 Department of Geosciences, University of Massachusetts, Amherst, Massachusetts U.S.A. Email: [email protected] ABSTRACT The taxonomy, phylogeny and biostratigraphy of the Oligocene Globanomalinidae (comprising the genera Pseudohastigerina and Turborotalia) is reviewed. Members of the Family have a macroperforate, nonspinose, smooth to weakly cancellate wall. Extinctions in the late Eocene left the Family restricted to just a few surviving forms which themselves became extinct in the early Oligocene. The group is very useful for biostratigraphy, with the extinctions of Pseudohastigerina naguewichiensis and Turborotalia ampliapertura providing the lowest two biozone boundaries of the Oligocene. The following species are recognized as valid: Pseudohastigerina micra (Cole), Pseudohastigerina naguewichiensis (Myatliuk), Turborotalia ampliapertura (Bolli), and Turborotalia increbescens (Bandy). INTRODUCTION The Globanomalinidae (as emended in this work) is an exclusively Paleogene Family with origins in the earliest Cenozoic. The first species, Globanomalina archeocompressa, evolved in the immediate aftermath of the Cretaceous/Paleogene mass extinction (Olsson and others, 1992) and was the root stock of a minor radiation of mostly smooth-walled macroperforate species with trochospiral ( Globanomalina and Turborotalia) or planispiral (Pseudohastigerina and Planoglobanomalina) tests. By the late Eocene, the Family was moderately diverse and abundant, especially the Turborotalia cerroazulensis group of species Cushman Foundation Special Publication No. 46 p. 403-414, 2018 (the evolution of which was studied by Pearson and Ezard, 2014), but it suffered in the extinctions of the Eocene-Oligocene transition. The Oligocene forms included herein are the last remnants of the Eocene diversification and the two groups (Pseudohastigerina and Turborotalia) are very divergent in morphology. The wall texture of Oligocene Turborotalia departs from the ancestral smooth condition and exhibits instead a pustulose to weakly cancellate structure. Their sequential extinction is vital for the biostratigraphic subdivision of the lower Oligocene (e.g., Bolli, 1957; Blow, 1979; Spezzaferri, 1994; Berggren and Pearson, 2005; Wade and others, 2011). A species-level range- chart and phylogeny is presented in Figure 14.1.
Transcript

Chapter 14

TAXONOMY, BIOSTRATIGRAPHY, AND PHYLOGENY OF OLIGOCENE GLOBANOMALINIDAE (PSEUDOHASTIGERINA AND TURBOROTALIA)

Paul N. Pearson1, Richard K. Olsson2, Silvia Spezzaferri3, and R. Mark Leckie4

1School of Earth and Ocean Sciences, Cardiff University, Main Building, Park Place, Cardiff, CF10 3AT, U.K. Email: [email protected]

2Department of Earth and Planetary Sciences, Busch Campus, Rutgers University, Piscataway, NJ 08854, U.S.A. Email: [email protected]

3Department of Geosciences, Earth Sciences, University of Fribourg, Ch du Musée 6, 1700 Fribourg, Switzerland. Email: [email protected]

4Department of Geosciences, University of Massachusetts, Amherst, Massachusetts U.S.A. Email: [email protected]

ABSTRACT

The taxonomy, phylogeny and biostratigraphy of the Oligocene Globanomalinidae (comprising the genera Pseudohastigerina and Turborotalia) is reviewed. Members of the Family have a macroperforate, nonspinose, smooth to weakly cancellate wall. Extinctions in the late Eocene left the Family restricted to just a few surviving forms which themselves became extinct in the early Oligocene.

The group is very useful for biostratigraphy, with the extinctions of Pseudohastigerina naguewichiensis and Turborotalia ampliapertura providing the lowest two biozone boundaries of the Oligocene. The following species are recognized as valid: Pseudohastigerina micra (Cole), Pseudohastigerina naguewichiensis (Myatliuk), Turborotalia ampliapertura (Bolli), and Turborotalia increbescens (Bandy).

INTRODUCTION

The Globanomalinidae (as emended in this work) is an exclusively Paleogene Family with origins in the earliest Cenozoic. The first species, Globanomalina archeocompressa, evolved in the immediate aftermath of the Cretaceous/Paleogene mass extinction (Olsson and others, 1992) and was the root stock of a minor radiation of mostly smooth-walled macroperforate species with trochospiral (Globanomalina and Turborotalia) or planispiral (Pseudohastigerina and Planoglobanomalina) tests. By the late Eocene, the Family was moderately diverse and abundant, especially the Turborotalia cerroazulensis group of species

Cushman Foundation Special Publication No. 46 p. 403-414, 2018

(the evolution of which was studied by Pearson and Ezard, 2014), but it suffered in the extinctions of the Eocene-Oligocene transition. The Oligocene forms included herein are the last remnants of the Eocene diversification and the two groups (Pseudohastigerina and Turborotalia) are very divergent in morphology. The wall texture of Oligocene Turborotalia departs from the ancestral smooth condition and exhibits instead a pustulose to weakly cancellate structure. Their sequential extinction is vital for the biostratigraphic subdivision of the lower Oligocene (e.g., Bolli, 1957; Blow, 1979; Spezzaferri, 1994; Berggren and Pearson, 2005; Wade and others, 2011). A species-level range-chart and phylogeny is presented in Figure 14.1.

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SYSTEMATIC TAXONOMY

Order FORAMINIFERIDA d’Orbigny, 1826Superfamily GLOBIGERINOIDEA Carpenter, Parker, and Jones, 1862

Family GLOBANOMALINIDAE Loeblich and Tappan, 1984, herein emended

Type Genus: Globanomalina Haque, 1956

DISCUSSION.— Loeblich and Tappan (1984; see also Loeblich and Tappan 1988) erected the Family Globanomalinidae to unite Globanomalina with forms ancestral to the Hantkeninidae (e.g. Clavigerinella) that were then considered to be directly related to each other. Subsequent research (Coxall and others, 2003) has disproved this link. In the Atlas of Eocene Planktonic Foraminifera, the Family Globanomalinidae was not recognized and the genera that are included in this work

were assigned to the Hedbergellidae (Olsson and Hemle-ben, 2006; Pearson and others, 2006a). Recent work on Cretaceous Hedbergella (especially wall microstructure studies) has revealed complexities in the phylogeny that casts significant doubt on a direct link between true Hedbergella from the Aptian (type species Anomalina lorneiana trochoidea Gandolfi, 1942) and later forms that have traditionally been included in Hedbergella, including the Maastrichtian ancestors of Paleocene Glo-banomalina (Huber and Leckie, 2011:61). Fundamental revisions to the Mesozoic suprageneric classification are likely to be implemented by the Mesozoic Plank-tonic Foraminifera Working Group (Huber and Leckie, 2011:61) which may include a restriction of Family Hedbergellidae (B.T. Huber, personal communication, 2011). For this reason the Family Globanomalinidae is resurrected and emended herein to include Globanom-alina and all its descendant forms (Pseudohastigerina, Planoglobanomalina, and Turborotalia).

Genus Pseudohastigerina Banner and Blow, 1959

TYPE SPECIES.— Nonion micrus Cole, 1927.

DISTINGUISHING FEATURES.— “The genus is characterized by a planispirally coiled, smooth-walled, normal perforate test, with an equatorial aperture which may vary from asymmetrical to symmetrical in position. The primary aperture may be singular or bipartite, arched openings bordered by a thin lip [centered about the axial periphery]” (Olsson and Hemleben, 2006:420).

DISCUSSION.— This genus was reviewed by Olsson and Hemleben (2006).

PHYLOGENETIC RELATIONSHIPS.— Pseudo-hastigerina evolved from Globanomalina luxorensis at the base of Zone E2 (Speijer and Samir, 1997; Berggren and Pearson, 2005; Olsson and Hemleben, 2006).

STRATIGRAPHIC RANGE.— Base of lower Eocene Zone E2 (Speijer and Samir, 1997; Molina and others, 1999; Berggren and Pearson, 2005) to the top of lower Oligocene Zone O1 (used as a zonal marker by Bolli, 1957; calibrated to Chron C12r by Miller and others, 1985; Biozone denoted O1 [last occurrence of P. naguewichiensis] by Berggren and Pearson, 2005).

GEOGRAPHIC DISTRIBUTION.—Global.

FIGURE 14.1. Stratigraphic ranges of the Oligocene Globanom-alinidae. Note that both increbescens and ampliapertura are now considered to have evolved from pomeroli in the middle Eocene; ampliapertura around the base of Zone E12 and increbescens around the middle of Zone E13 (see text for discussion).

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Pseudohastigerina micra (Cole, 1927)

(Note: a more extensive synonymy list was provided by Olsson and Hemleben, 2006. Here we focus on

Oligocene occurrences.)

Nonion micrus Cole, 1927:22, pl. 5, fig. 12 [middle Eocene Zone E11, Guayabal Fm., Tampico, Mexico].

Pseudohastigerina micra (Cole).—Berggren and others, 1967:275, text-fig. 9 [middle Eocene Zone P12, type locality, Guayabal Fm., Mexico].—Leckie and others, 1993:125, pl. 6, figs. 14-17 [lower Oligocene Zone P18, ODP Hole 628A, western North Atlantic Ocean].—Olsson and Hemleben, 2006:422-424, pl. 14.3, figs. 11, 12 [SEM of holotype], figs. 13, 14 [SEM of holotype of Nonion danvillensis Howe and Wallace, 1932, Zone E15/16, Jackson Fm., Danville Landing, Louisiana], fig. 15 [middle Eocene Zone E8, Tanzania Drilling Project Site 2/18/1, 20-26 cm, Kilwa, Tanzania], figs. 16, 19 [upper Eocene Zone E15/16, Jackson Fm., Danville Landing, Louisiana], figs. 17, 18, 22 [upper Eocene, Atlantic City borehole, New Jersey, ODP 150X: 1338.0-.1 feet], figs. 20, 21, 24 [middle Eocene Zone E9, Tanzania Drilling Project Site 2/9/CC, Kilwa, Tanzania].—Wade and Olsson, 2009, pl. 1, figs. f-m [upper Eocene Zone E15/16, TDP Site 12, Stakishari, Tanzania].—Pearson and Wade, 2015, fig. 26.1, 26.3, 26.4, 26.6 [upper Eocene Zone E15/16, TDP Site 12, Stakishari, Tanzania], fig. 26.2, 26.5, 26.7 [upper Eocene Zone E15/16, TDP Site 17, Stakishari, Tanzania].

Pseudohastigerina aff. P. micra (Cole).—Leckie and others, 1993, pl. 6, fig. 20 [lower Oligocene Zone P18, ODP Hole 628A, western North Atlantic Ocean].

DESCRIPTION.Type of wall: Smooth, normal perforate.Test morphology: Test planispiral, compressed,

tightly coiled, involute, circular to oval in outline; in spiral view 6-8 chambers in ultimate whorl, increasing slowly in size, sutures slightly depressed, straight to gently curved between ultimate chambers, chambers slightly inflated, umbilicus small, circular in shape, generally only apertural lip of ultimate chamber visible; in edge view, primary aperture equatorial, symmetric, a circular high arch bordered by a narrow lip, bipartite apertures sometimes present on ultimate chamber, test compressed with a subrounded to subacute periphery (modified from Olsson and Hemleben, 2006).

Size: maximum diameter of holotype 0.17 mm, thickness 0.09 mm.

DISTINGUISHING FEATURES.— This species is distinguished from P. naguewichiensis by its more rapid rate of chamber enlargement, more compressed periphery, and generally more involute coiling. In the Eocene it is also distinguished by its generally larger size, but both species are always small in the Oligocene, making discrimination more difficult (see, for example, Leckie and others, 1993:125).

DISCUSSION.— Olsson and Hemleben (2006) provided a full discussion and illustration of this species. Pseudohastigerina micra is common in the upper Eocene but it declined dramatically in size and abundance at the Eocene/Oligocene boundary (Cordey and others, 1970; Keller, 1985, Nocchi and others, 1986; Miller and others, 2008; Wade and Pearson, 2008). The disappearance of large-sized P. micra can be used to correlate the Eocene/Oligocene boundary proper, especially when hantkeninids are rare or absent (Miller and others, 2008; Wade and Pearson, 2008).

PHYLOGENETIC RELATIONSHIPS.— Pseudo-hastigerina micra evolved from Pseudohastigerina wilcoxensis (Olsson and Hemleben, 2006).

STRATIGRAPHIC RANGE.— Zone E7 (Olsson and Hemleben, 2006) to Zone O1 (Keller, 1985; Nocchi and others, 1986; Leckie and others, 1993; Pearson and Chaisson, 1997; Wade and Pearson, 2008). The species is typically rare in Zone O1 and it is unclear how far through the zone the range extends. The highest occurrence we have confirmed is from the middle of Zone O1 (Leckie and others, 1993, pl. 6, fig. 20; recorded as Pseudohastigerina aff. P. micra).

TYPE LEVEL.— Middle Eocene Zone E11, Guayabal Fm., Mexico.

GEOGRAPHIC DISTRIBUTION.— Global in low to high latitudes.

STABLE ISOTOPE PALEOBIOLOGY.— This species usually registers among the most negative δ18O values of assemblages, indicating a shallow water, mixed-layer habitat. Carbon isotopes are strongly depleted as is common in small-sized non-symbiotic species (Poore and Matthews, 1984; Boersma and others, 1987; Pearson and others, 2001).

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REPOSITORY.— Holotype deposited at the Paleontological Research Institution, Ithaca, New York. Paratype (USNM CC243208) deposited at the Smithsonian Museum of Natural History, Washington, D.C.

Pseudohastigerina naguewichiensis (Myatliuk, 1950)

Plate 14.1, Figures 1-16

(Note: A more extensive synonymy list was presented by Olsson and Hemleben, 2006)

Globigerinella naguewichiensis Myatliuk, 1950:281, pl. 4, figs. 4 a, b [Oligocene, Chechva River, Lopanetskie layer, western Ukraine].—Subbotina, 1953:124, pl.13, fig. 18a, b [re-illustration of holotype].

Pseudohastigerina naguewichiensis (Myatliuk).—Blow, 1969:377, pl. 53, figs. 2, 3 [Oligocene Zone P18, Red Bluff Clay type locality, Mississippi (originally stated as Alabama)].—Olsson and Hemleben, 2006:424-426, pl. 14.3, figs. 1, 2 [holotype of P. barbadoensis Blow [Oligocene Zone P19, Cipero Fm., Trinidad], figs. 3, 4, 6-8 [Zone E15/16, Shubuta Clay Member, Yazoo Fm., Chicksawhay River, Wayne County, Mississippi], figs. 5, 9 [Zone O1, TDP Site 11, Stakishari, Tanzania], fig. 10 [upper Eocene, Atlantic City borehole, New Jersey, ODP 150X, 1338.0-.1 feet].—Wade and Olsson, 2009, pl. 1, figs. a, b [lower Oligocene Zone O1, St. Stephen’s Quarry, Alabama], fig. c [upper Eocene Zone E16, St. Stephen’s Quarry, Alabama], figs. d, e [lower Oligocene Zone O1, TDP Site 11].—Pearson and Wade, 2015:23, fig. 26.8 [lower Oligocene Zone O1, TDP Site 12, Stakishari, Tanzania], fig. 26.9, 26.11 (reproduced from Olsson and Hemleben, 2006, pl. 14.3, figs. 9, 5), 26.10 [lower Oligocene Zone O1, TDP Site 17, Stakishari, Tanzania].

Pseudohastigerina cf. P. naguewichiensis (Myatliuk).—Leckie and others, 1993:125, pl. 6, figs. 18, 19 [lower Oligocene Zone P18, ODP Hole 628A, western North Atlantic Ocean, and Hole 803D, western equatorial Pacific Ocean].

Pseudohastigerina barbadoensis Blow, 1969:409, pl. 53, figs. 7-9 [Oligocene Zone P19, Cipero Fm., Trinidad (figs. 7, 8 holotype; fig. 9 paratype)], pl. 54, figs. 1-3, paratypes [Oligocene Zone P19, Cipero Fm., Trinidad].

Plate 14.1 Pseudohastigerina naguewichiensis (Myatliuk, 1950)

1-2 (holotype; reproduced from Myatliuk, 1950, pl. 4, fig. 4a-b; Oligocene, Kosmach series, Chechva River, North Carpathians); 3, 4 (holo-type of Pseudohastigerina barbadoensis Blow, 1969, pl. 53, figs. 7, 8; Oligocene Zone P19/20, Cipero Fm., Trinidad); 5, Zone O1, Sample TDP 11/25/3, 10-20 cm, Pande Fm., Tanzania (reproduced from Olsson and Hemleben, 2006, pl. 14.3, fig. 5), 6, Zone O1, Sample TDP 17/11/1, 0-10 cm, Pande Fm., Tanzania; 7, 8 (same specimen), Zone O1, Sample TDP 11/19/1, Pande Fm., Tanzania; 9-12, Zone O1, AGS 66, 9A-1A, Shubuta Fm., Alabama; 13, Zone E16, Shubuta Fm., 158 ft., St. Stephen’s Quarry, Alabama; 14-16 (same specimen), Zone O1, ODP 706A/3H/4, 93-95 cm, Mascarene Plateau, tropical Indian Ocean. Scale bar: 1-16 = 50 µm.

DESCRIPTION.Type of wall: “Smooth, normal perforate; in

adult stage wall becomes thickened (gametogenetic calcite?) and openings to pores are enlarged, giving appearance of a coarsely perforate wall” (Olsson and Hemleben, 2006:424). Periphery may be imperforate.

Test morphology: Test small, planispiral, compressed, tightly coiled, evolute, circular to oval in outline, slightly lobulate, chambers weakly inflated; in umbilical view 6-9 chambers in ultimate whorl, increasing very slowly in size, circular in outline, sutures slightly depressed, straight to slightly curved, umbilicus circular in shape, inner coil of chambers partly visible; in edge view chambers globular, slightly oval to nearly circular in outline, primary aperture a moderately high arch bordered by a thickened prominent lip, test compressed with a rounded to subrounded periphery, peripheral margin perforate, some pores closed off by thickened wall (modified from Olsson and Hemleben, 2006).

Size: Maximum diameter of holotype 0.20 mm, thickness 0.09 mm.

DISTINGUISHING FEATURES.— Pseudohastigerina naguewichiensis is distinguished from P. micra by the greater inflated chambers, more evolute coiling, and the slowly increasing chamber size, which causes a more circular test outline. The margin is typically more rounded than in P. micra. It has a tendency to be heavily encrusted especially on the periphery, with pore pits in the adult stage.

DISCUSSION.— This species was discussed by Olsson and Hemleben (2006) who followed Toumarkine and Luterbacher (1985) in regarding P. barbadoensis Blow as a junior synonym. The type specimen of P. naguewichiensis is lost; the illustration reproduced on Plate 14.1, Figs. 1-2, shows quite clearly a planispiral evolute form with relatively straight sutures, a lobate outline and only six chambers in the final whorl. The holotype of P. barbadoensis, reproduced on Plate 14.1, Figs. 3-4, is also evolute but is larger, less lobate and has

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Plate 14.1 Pseudohastigerina naguewichiensis (Myatliuk, 1950)

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Pearson, Olsson, Spezzaferri, and Leckie

Test morphology: “Trochospiral, varying from globular and inflated to strongly compressed and biconvex; flattened on spiral side; primary aperture umbilical-extraumbilical, usually a high arch with bordering lip; imperforate band or keel present in some species” (Pearson and others, 2006b:438).

DISCUSSION.— This genus was reviewed by Pearson and others (2006b). The two species that survive into the Oligocene are quite similar and form a spectrum of morphology from the more ‘globigerinid’ T. ampliapertura to the more ‘globorotaliid’ T. increbescens. Of the two, T. ampliapertura is by far the most abundant (e.g., Leckie and others, 1993; Wade and Pearson, 2008).

STRATIGRAPHIC RANGE.— Lower Eocene to Oligocene.

Turborotalia ampliapertura (Bolli, 1957)

Plate 14.2, Figures 1-16

Globigerina ampliapertura Bolli, 1957:108, pl. 22, figs. 4a-6c [lower Oligocene Globigerina ampliapertura Zone, Cipero Fm., Trinidad].

“Turborotalia” ampliapertura (Bolli).— Leckie and others, 1993:125-126, pl. 4, figs. 3-8 [lower Oligocene Zone P18, ODP Hole 628A, western North Atlantic Ocean].

Turborotalia ampliapertura (Bolli).—Pearson and others, 2006b:441-442, pl. 15.2, figs. 1-3 [SEMs of holotype], figs. 4-6, 8 [lower Oligocene Globigerina ampliapertura Zone, Cipero Fm., Trinidad], figs. 7, 17-20 [Zone E15/16, Shubuta Clay, Wayne County, Mississippi], figs. 9-11 [SEM of holotype of Globigerina pseudoampliapertura Blow and Banner, Zone E16, Sample FCRM 1923, Madingura (= Namadingura) River, Lindi, Tanzania], figs. 12-16 [topotypes of Globigerina pseudoampliapertura Blow, Zone E16, Sample FCRM 1923, Madingura (= Namadingura) River, Lindi, Tanzania].—Pearson and Wade, 2015, fig. 27.1-27.6 [lower Oligocene Zone O1, TDP Site 17, Stakishari, Tanzania].

Globigerina pseudoampliapertura Blow and Banner, 1962:95, pl. 12, figs. a-c, pl. 17, figs. a, e [upper Eocene, Sample FCRM 1923, Madingura (= Namadingura) River, Lindi, Tanzania].

Globigerina kondoi Todd, 1970:A16, pl. 7, fig. 2a-c [upper Eocene, southeastern Eua Island, Tonga].

?Turborotalia pseudoampliapertura nukhulensis Haggag and Luterbacher, 1995, pl. 2, figs. 5-9 [uppermost part of middle Eocene Morozovella lehneri Zone, Wadi Nukhul,

more curved sutures. However, like P. naguewichiensis, P. barbadoensis has more inflated chambers, a more rounded periphery, and a more coarsely perforate test than P. micra. Together these specimens illustrate much of the range of variation shown by this species.This synonymy of barbadoensis with naguewichiensis was also discussed by Pearson and Wade (2015).

PHYLOGENETIC RELATIONSHIPS.— Pseudo-hastigerina naguewichiensis evolved from P. micra (Olsson and Hemleben, 2006).

STRATIGRAPHIC RANGE.— Zone E14 (Cotton and others, 2017) to O1 (Bolli, 1957; Leckie and others, 1993; Pearson and Chaisson, 1997; Berggren and Pearson, 2005; see also comments under Genus Pseudohastigerina, above).

TYPE LEVEL.— Originally described as ‘Oligocene’, probably lower Oligocene Zone O1, Kosmach series, North Carpathians, Ukraine.

GEOGRAPHIC DISTRIBUTION.— Global in low to high latitudes.

STABLE ISOTOPE PALEOBIOLOGY.— Poore and Matthews (1984) and Wade and Pearson (2008) recorded relatively negative oxygen isotopes for this species indicating a surface water, mixed-layer habitat as in other Pseudohastigerina.

REPOSITORY.— Holotype (No. 2321) deposited in the VNIGRI collection, St. Petersburg, Russia.

Genus Turborotalia Cushman and Bermúdez, 1949

Globorotalia (Turborotalia) Cushman and Bermúdez, 1949:42Turborotalia Bykova and others, 1959

TYPE SPECIES.— Globorotalia centralis Cushman and Bermúdez, 1937 (=junior subjective synonym of Globigerina cerro-azulensis Cole, 1928; see Pearson and others, 2006b).

DESCRIPTION.Type of wall: Smooth to pustulose, sometimes

weakly cancellate; normal size pores.

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ampliapertura were named, nukhulensis and sinaiensis. In our view the latter falls within the normal range of variation of ampliapertura. The former is somewhat intermediate between pomeroli and ampliapertura and is here included in questionable synonymy with ampliapertura. The study of Haggag and Luterbacher (1995) demonstrates that ampliapertura evolved around the base of Zone E12, that is, during the middle Eocene climate optimum. Its ancestor was evidently pomeroli, not increbescens as suggested by Blow and Banner (1962) and Pearson and others (2006b).

PHYLOGENETIC RELATIONSHIPS.— Evolved from Turborotalia pomeroli in the middle Eocene (Haggag and Luterbacher, 1995). It became extinct in the Oligocene without leaving any descendants, bringing to an end the stratigraphic range of the Family Globanomalinidae.

STRATIGRAPHIC RANGE.— This form ranges from around the base of middle Eocene Zone E12 (Haggag and Luterbacher, 1995, fig. 2) to lower Oligocene Zone O2 (used as a zone marker by Bolli, 1957; calibrated to Chron C11r by Leckie and others, 1993; zone denoted O2 by Berggren and Pearson, 2005).

TYPE LEVEL.— Lower Oligocene Zone O1-O2 (Globigerina ampliapertura Zone), Cipero Fm., Trinidad.

GEOGRAPHIC DISTRIBUTION.— Cosmopolitan, occurring across a broad range of latitudes. The species peaks in abundance during the Eocene-Oligocene turnover and after the extinction of other Turborotalia suggesting it is an opportunistic species (Wade and Pearson, 2008).

STABLE ISOTOPE PALEOBIOLOGY.— Poore and Matthews (1984) recorded consistently negative δ18O values for this species indicating a shallow water, mixed-layer habitat. Wade and Pearson (2008) found it consistently offset by about +0.5 per mil from co-occurring Pseudohastigerina micra, suggesting a near surface habitat but perhaps with a component of gametogenic calcite added at depth.

REPOSITORY.— Holotype (USNM P5603) deposited at the Smithsonian Museum of Natural History, Washington, D.C.

Egypt].Turborotalia pseudoampliapertura sinaiensis Haggag and

Luterbacher, 1995, pl. 3, figs. 1-4 [upper part of middle Eocene Truncorotaloides rohri Zone, Wadi Nukhul, Egypt].

DESCRIPTION. Type of wall: Smooth, to weakly cancellate, frequently densely pustulose; tendency to defoliate. Test morphology: Moderately high trochospiral, compact globular test with 3½-4 chambers in final whorl. Chambers inflated, appressed and embracing, increasing moderately in size. Outline lobulate or rounded in edge view. Spiral sutures slightly curved, depressed. Aperture a high arch, sometimes asymmetrical but approaching circular in some specimens, in umbilical-extraumbilical position; may have imperforate rim. Umbilicus small. Umbilical sutures moderately curved, depressed. The coiling is somewhat irregular and there is considerable variation between specimens in the shape and size of the aperture (modified from Pearson and others, 2006b). Size: Holotype length 0.49 mm, width 0.29 mm.

DISTINGUISHING FEATURES.— Turborotalia ampliapertura is distinguished from T. increbescens by a more umbilically centered aperture and more globular chamber shape (Pearson and others, 2006b). It is common for populations to show some intergradation between these species. Specimens on Plate 14.2 have been selected to illustrate the range of variation. Some specimens can be similar in shape to Dentoglobigerina taci Pearson and Wade but are distinguished by having less incised sutures and a different wall texture. DISCUSSION.— Turborotalia ampliapertura is a useful zone fossil (Bolli, 1957). Pearson and others (2006b) included Globigerina pseudoampliapertura Blow and Banner in synonymy. The evolutionary origin of ampliapertura has been somewhat problematic, with the first appearance being given as upper Eocene Zone E15 (Blow and Banner, 1962; Pearson and others, 2006b). However the most detailed information is provided by Haggag and Luterbacher (1995) who studied the evolutionary transition between Turborotalia pomeroli and Turborotalia ampliapertura (given as pseudoampliapertura) in the middle Eocene of Egypt. In that study (which was overlooked by Pearson and others, 2006b) two subspecies of pseudo-

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PLATE 14.2 Turborotalia ampliapertura (Bolli, 1957)

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Chapter 14 - Pseudohastigerina and Turborotalia

Turborotalia increbescens (Bandy, 1949)

Plate 14.3, Figures 1-15

Globigerina increbescens Bandy, 1949:120, pl. 23, figs. 3a-c [upper Eocene, Little Stave Creek, Alabama].—Snyder and Waters, 1985:460, pl. 2, fig. 12-14 [lower Oligocene Zone P20, DSDP Hole 549A, Goban Spur, North Atlantic Ocean].

Globorotalia (Turborotalia) increbescens (Bandy).—Blow and Banner, 1962:118, pl. 13, figs. T-V, pl. 17, figs. D, K [upper Eocene, Sample FCRM. 1650, Lindi, Tanzania].—Blow, 1969:349, pl. 36, figs. 5, 6 [upper Eocene, Lindi, Tanzania].

Turborotalia increbescens (Bandy).—Leckie and others, 1993:126, pl. 4, fig. 1 [lower Oligocene Zone P18, ODP Hole 803D, Ontong Java Plateau, western equatorial Pacific Ocean], fig. 2 [lower Oligocene Zone P18, ODP Hole 628A, western North Atlantic Ocean].—Nishi and Chaproniere, 1994:260, pl. 4, figs. 7-9 [lower Oligocene Zone P18, ODP Hole 841B, Tonga Trench, South Pacific Ocean].—Pearson and others, 2006b:453-454, pl. 15.6, figs. 1-3 [SEMs of holotype], figs. 4-14 [SEMs of paratypes], fig. 15 [Zone E14, upper Eocene, Kitunda, near Lindi, Tanzania].—Pearson and Wade, 2015, fig. 27.7a-b [lower Oligocene Zone O1, TDP Site 12, Stakishari, Tanzania], fig. 27.8a-b [lower Oligocene Zone O1, TDP Site 17, Stakishari, Tanzania].

DESCRIPTION. Type of wall: High porosity, weakly cancellate and coarsely pustulose, often smooth on final chamber; like other Turborotalia it has a tendency to defoliate. Test morphology: “Moderate to high trochospiral, globorotaliform inner whorl, compact and rounded test with 3½-4 chambers in the final whorl. Chambers appressed and embracing, radially compressed and increasing moderately in size. Final chamber commonly radially flattened; in edge view, showing rounded periphery with final chamber arching towards umbilicus. Dorsal sutures curved, depressed. Aperture a very broad arch in an intra-extraumbilical position, often irregular and elongate. An imperforate or pustulose lip is usually visible. Umbilicus usually narrow. Ventral sutures slightly curved, depressed. Usually a strong tendency for sinistral coiling” (Pearson and others, 2006b:454). Size: Holotype length 0.51 mm, breadth 0.47 mm.

DISTINGUISHING FEATURES.— “Turborotalia increbescens is distinguished from both T. pomeroli and T. cerroazulensis by its more compact morphology and from T. cerroazulensis by having a rounder periphery and more globular test shape” (Pearson and others, 2006b:454). It is distinguished from T. ampliapertura by its longer inner whorl, distinctly more extraumbilical position of the aperture and less globular chamber shape, which gives the test a more ‘globorotaliid’ appearance.

DISCUSSION.— This species was discussed by Pearson and others (2006b). Like T. ampliapertura it survived the extinction of the rest of the genus in the uppermost Eocene. It is rarer and more sporadic than T. ampliapertura in the Oligocene (e.g., Leckie and others, 1993).

PHYLOGENETIC RELATIONSHIPS.— Evolved from Turborotalia pomeroli in the middle Eocene (Blow and Banner, 1962; Toumarkine and Bolli, 1970). The species is no longer considered the likely ancestor of ampliapertura (see discussion under that species).

STRATIGRAPHIC RANGE.— Middle Eocene Zone E13 (Postuma, 1971) to lower part of lower Oligocene Zone O2 (Pearson and Chaisson, 1997). Blow (1979, fig. 53, p. 311) and Leckie and others (1993, p. 118) recorded the highest occurrence as slightly below the topmost Pseudohastigerina and hence in the top of Zone O1; Pearson and Chaisson (1997) recorded it extending to just within (the lowermost of 8 studied samples) Zone O2.

TYPE LEVEL.— Probably E16, upper part of Jackson Fm., upper Eocene, Little Stave Creek, Alabama.

GEOGRAPHIC DISTRIBUTION.— Cosmopolitan, more common at low latitudes (Leckie and others, 1993).

STABLE ISOTOPE PALEOBIOLOGY.— Recorded by Boersma and others (1987) as having relatively negative δ18O, possibly indicating a shallow-water habitat.

REPOSITORY.— Holotype deposited in the micropaleontological collections at Indiana University.

Plate 14.2 Turborotalia ampliapertura (Bolli, 1957)

1-3 (holotype, USNM P5603; reproduced from Pearson and others, 2006b, pl. 15.2, figs. 1-3); 4-8, Zone O1, Sample TDP 17/11/1, 0-10 cm, Pande Fm., Tanzania; 9-12, Zone O1, Shubuta Fm., Alabama; 13-16, Zone O2, Cipero Fm., Trinidad. Scale bar: 1-16 = 100 µm.

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Plate 14.3 Turborotalia increbescens (Bandy, 1949)

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Chapter 14 - Pseudohastigerina and Turborotalia

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Plate 14.3 Turborotalia increbescens (Bandy, 1949)

1-3 (holotype, PLIU 4904; reproduced from Pearson and others, 2006b, pl. 15.6, figs. 1-3); 4-11, Zone O1, Sample TDP 17/11/1, 0-10 cm, Pande Fm., Tanzania; 12-15, Zone O1, Shubuta Fm., Alabama. Scale bars: 1-10, 12-15 = 100 µm; 11 = 10 µm.

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Citation

Pearson, P.N., Olsson, R.K., Spezzaferri, S., and Leckie, R.M., 2018, Taxonomy, biostratigraphy, and phylogeny of Oligocene Globanomalinidae (Pseudohastigerina and Turborotalia) in Wade, B.S., Olsson, R.K., Pearson, P.N., Huber, B.T. and Berggren, W.A. (eds.), Atlas of Oligocene Planktonic Foraminifera, Cushman Foundation of Foramin-iferal Research, Special Publication, No. 46, p. 403-414.


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