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Taxonomy, biostratigraphy, and phylogeny of Eocene Morozovella

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CUSHMAN FOUNDATION SPECIAL PUBLICATION NO. 41 P. 343-376, 2006
Chapter 11
TAXONOMY, BIOSTRATIGRAPHY, AND PHYLOGENY OF EOCENE
MOROZOVELLA
WILLIAM A. BERGGREN1,2 AND PAUL N. PEARSON3
1Department of Geology and Geophysics, Woods Hole Oceanographic Institution, Woods Hole, MA 02543,
U.S.A. Email. wberggren@whoi.edu
2Department of Geological Sciences, Rutgers University, Busch Campus, Piscataway, NJ 08854=8066, U.S.A.
3School of Earth, Ocean and Planetary Sciences, Cardiff University, Main Building, Park Place, Cardiff CF10
3YE, U.K. Email: pearsonp@cardiff.ac.uk
ABSTRACT
Following their origin in the Paleocene, morozovellids
underwent a renewed radiation in the early Eocene
and split into two main groups: a low anguloconical
group represented by the M. aequa-subbotina-gracilis-
formosa morphotypes and a high angloconical group
represented by the lensiformis-aragonensis-causcasica
morphotypes. The morozovellids became extinct in
the early middle Eocene with the LAD of M.
aragonensis, which marks the top of Zone E9. In this
chapter we discuss the taxonomy, biostratigraphy and
phylogenetic relations of eight species of Morozovella
which are restricted to the Eocene: M. allisonensis,
M. aragonensis, M. caucasica, M. crater, M. edgari,
M. formosa, M. gracilis, and M. marginodentata. Eight
other species (M. acuta, M. aequa, M. apanthesma,
M. lensiformis, M. occlusa, M. pasionensis, M.
subbotinae, and M. velascoensis) range into basal
Eocene strata and were treated in the Atlas of
Paleocene Planktonic Foraminifera (Olsson and
others, 1999). Several middle Eocene taxa heretofore
placed in Morozovella (bandyi, crassata, coronata,
lehneri) are here excluded and assigned to the
homeomorphic genus Morozovelloides (Pearson and
Berggren; see Chapter 10, this volume).
INTRODUCTION
In their initial Paleocene radiation, the
morozovellids split into two main groups: (1) the M.
angulata-M. velascoensis group characterized by the
development of muricate adumbilical ridges, a
muricocarina, and the absence of muricae on parts of the
chamber surfaces; and (2) the M. apanthesma-M.
subbotinae group whose members were initially unkeeled
and entirely covered with fine, thin muricae.
In the early Eocene, the velascoensis group
became extinct, and the surviving morozovellids
diversified further along two distinct morphogenetic
trends: (1) a low anguloconical group represented by
the M. aequa-subbotina-gracilis-formosa group; and (2)
a high angloconical group represented by the M.
lensiformis-aragonensis-caucasica group. This latter
group produced morphologies that are highly
convergent on the previous velascoensis group.
Morozovella aragonensis (LAD at the top of Zone E9)
is the terminal member of the genus Morozovella as
delimited here. The species level range-chart and
phylogeny is presented in Figure 11.1.
BERGGREN AND PEARSON
344
FIGURE 11.1. Stratigraphic ranges and inferred phylogenetic relationships of Eocene species of Morozovella discussed in this chapter.
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CHAPTER 11 - MOROZOVELLA
345
SYSTEMATIC TAXONOMY
Order FORAMINIFERIDA Eichwald, 1830
Superfamily GLOBIGERINACEAE Carpenter,
Parker and Jones, 1862
Family TRUNCOROTALOIDIDAE Loeblich and
Tappan, 1961
Genus Morozovella McGowran in Luterbacher,
1964
TYPE SPECIES.— Pulvinulina velascoensis Cushman,
1925
DESCRIPTION.
Type of wall: Normal perforate (pores cylindrical
on at least part of the later chambers); some forms with
weakly cancellate early chambers (a feature inherited
from their praemuricate ancestry); adult chambers
strongly pustulose (muricate) on parts of the spire and
umbilicus.
Test morphology: Test low trochospiral, lobulate
outline; strongly anguloconical chambers throughout
ontogeny; peripheral margin usually bears distinct
muricocarina which passes continuously across at least
one intercameral suture (absent or rudimentary in
apanthesma); primary aperture interiomarginal,
umbilical-extraumbilical; secondary apertures may be
present on spiral side but typically lack apertural lips.
DISTINGUISHING FEATURES.— Genus distin-
guished by muricate, anguloconical test and peripheral
muricocarina (in nearly all taxa) of variable breadth and
strength.
DISCUSSION.— The genus Morozovella was originally
considered a subgenus of Truncorotaloides Brönnimann
and Bermúdez, 1953 by McGowran, in Luterbacher
(1964) based on the presence of “conical shaped
chambers”, the “thickened and keeled” or “subangular
to angular” peripheral margin. Blow (1979, p. 972)
emended the diagnosis of the genus Morozovella to
include forms in which the “subacute to acutely angular
peripheral test margin bears a muricocarina, at least
continuous from one chamber to the next successively
added chamber in some part of the last convolution of
chambers within the trochospire of the test”. A careful
reading of Blow (1979, p. 320, 396, 400, 972) suggests
that the main difference separating Paleogene
Morozovella and Neogene Globorotalia s.s. is the
presence of muricae and the nature of the peripheral
muricocarina in the Paleogene forms and the true
peripheral keel (formed from the lateral, marginal
compression of the primary wall) in the Neogene forms.
The morozovellids split into two major clades
early in their evolution: (1) the M. angulata-velascoensis
group characterized by muricate adumbilical ridges, a
strong muricocarina, and the absence of muricae on parts
of the chamber surfaces; and (2) the M. apanthesma-M.
subbotinae group whose members are initially unkeeled
and entirely covered with fine, thin muricae. Clade 1
appears to terminate in the early Eocene with the
sequential extinction of the Morozovella acuta-
velascoensis plexus and, finally, M. edgari. Clade 2
undergoes a demonstrable radiation in the early Eocene
(producing, among others, lensiformis, crater,
aragonensis, caucasica) and terminated in Zone E9 with
the extinction of M. aragonensis.
Morozovella has a stratigraphic range from the
mid-Paleocene (Zone P3: M. angulata) to middle Eocene
(Zone E9: M. aragonensis).
Morozovella acuta (Toulmin, 1941)
DISCUSSION.— This taxon was discussed in the Atlas
of Paleocene Planktonic Foraminifera (Olsson and
others, 1999).
STRATIGRAPHIC RANGE.— Zone P4b to Zone E2
(top).
Morozovella aequa (Cushman and Renz, 1942)
PLATE 11.1, FIGURES 1-8
DISCUSSION.—This taxon was discussed in the Atlas
of Paleocene Planktonic Foraminifera (Olsson and
others, 1999).
STRATIGRAPHIC RANGE.— Zone P4c to Zone E5
Morozovella allisonensis Kelly, Bralower, and
Zachos, 1998
PLATE 11.2, FIGURES 4-16
(Pl. 11.2, Figs. 4-8, 12: SEMs of holotype and
paratype of Morozovella allisonensis Kelly, Bralower,
and Zachos)
BERGGREN AND PEARSON
346
Morozovella allisonensis (nomen nudum) Kelly Bralower,
Zachos, Premoli Silva, and Thomas, 1996:424, figs. 2-7a
to 2-7b [Zone P5 (from a part equivalent to Zone E1 in
this work, from within carbon isotope excursion of
Paleocene-Eocene Thermal Maximum), ODP Hole 865C
Allison Guyot, equatorial Pacific Ocean].
Morozovella allisonensis Kelly, Bralower, and Zachos,
1998:158, figs. 5A [holotype], 5B [paratype]; figs. 8D-G
[Zone P5; (from a part equivalent to Zone E1 in this work,
from within carbon isotope excursion of Paleocene-Eocene
Thermal Maximum), ODP Hole 865C Allison Guyot,
equatorial Pacific Ocean].
DESCRIPTION.
Type of wall: Normal perforate, muricate,
nonspinose.
Test morphology: Low to medium trochospiral
test, subcircular, weakly to non-lobulate peripheral
outline, chambers triangular on umbilical side,
trapezoidal to subquadrate on spiral side as a function
of degree of curvature of intercameral sutures; in
umbilical view primary aperture low arch to subcircular,
umbilical to extraumbilical in position extending to
peripheral margin; typically 5-9 chambers in the final
whorl; sutures depressed, straight to slightly curved,
radial; umbilicus generally small, deep varying with
tightness of coiling; chambers with unornamented
rounded umbilical shoulders, slight tendency to be
umbilically inclined as a function of axial compression;
on the spiral side approximately 12-17 chambers
arranged in about 3 whorls; chambers flattened to
moderately inflated; gradual increase in chamber size
throughout; sutures muricate, strongly to weakly curved,
varying from raised to slightly depressed; in edge view
umbilico-convex to planar; spiral side flat to slightly
convex; test periphery subrounded to acute; weakly
muricate keel.
Size: Holotype maximum diameter:
approximately 0.35 to 0.40 mm.
DISTINGUISHING FEATURES.— This taxon is
characterized by varying degrees of axial compression;
it is biconvex to mildly planoconvex in edge view;
umbilical tips of chambers are relatively rounded and
unornamented.
DISCUSSION.— This taxon exhibits a wide range of
morphological plasticity that is a function of degree of
axial compression. We have observed the relatively high
conical morphotypes intermediate between the typical
end members of the velascoensis-allisonensis group
within the CIE in the Dababiya and Qreiya sections in
Egypt.
PHYLOGENETIC RELATIONSHIPS.— This
ephemeral taxon intergrades with more ornate
morphotypes of its ancestor, M. velascoensis, and has
no descendants.
STRATIGRAPHIC RANGE.— Zone E1; restricted to
Carbon Isotope Excursion (CIE) interval of Paleocene-
Eocene Thermal Maximum (PETM).
GEOGRAPHIC DISTRIBUTION.— This taxon is most
abundant in warm-water, (sub)tropical regions; common
in central-equatorial Pacific Ocean (ODP Site 865) and
in northern (Spain) and southern (Egypt) Tethyan
deposits; rare occurrences recorded on Blake Ridge of
North Atlantic (ODP Site 1051).
STABLE ISOTOPE PALEOBIOLOGY.— Inferred
depth-habitat varies over time; initially occupied deeper
portions of mixed-layer to upper thermocline and
subsequently inhabited shallow mixed-layer; carbon
isotope signature exhibits strong covariance with shell
size, while oxygen isotope composition displays a
modest negative correlation with shell size (overall stable
Plate 11.1, 1-8, Morozovella aequa (Cushman and Renz, 1942); 9-16, Morozovella subbotinae (Morozova, 1929)
Morozovella aequa 1, 2, 4, 5, 7, Zone E1, Bass River Borehole, New Jersey, ODP 174AX: 1145.0-1162.0 feet; 3, 6 (Blow, 1979,
pl. 96, figs. 2, 5, 6), Zone P5, Sample FCRM 1670, Lindi area, Tanzania, 8 (Morozovella aequa tholiformis Blow, 1979, pl. 102, fig. 8),
Zone E3/4, DSDP Hole 20C/63, 76-78 cm, central South Atlantic Ocean. Morozovella subbotinae 9-11 (Globorotalia rex Martin, 1943,
paratype, USNM 559452), Zone E3/4, Lodo Fm., Fresno Co., California; 12 (Blow, 1979, pl. 102, figs. 8, 12), Zone P6, DSDP Hole 20C/
63, 76-78 cm, South Atlantic Ocean; 13, 15 (Blow, 1979, pl. 111, figs. 7, 8), Zone E5, Moogli Mudstones, Kagua, Papua; 14, 16 (Blow,
1979, pl. 127, figs. 6, 7), Zone E5, DSDP Site 47/2/8, 71-73 cm, northwest Pacific Ocean. Scale bar: 1-16 = 100 µm.
CHAPTER 11 - MOROZOVELLA
347
PLATE 11.1 Morozovella aequa (Cushman and Renz, 1942), Morozovella subbotinae (Morozova, 1929)
BERGGREN AND PEARSON
348
PLATE 11.2 Morozovella velascoensis (Cushman, 1925), Morozovella allisonensis (Kelly, Bralower and Zachos, 1998)
CHAPTER 11 - MOROZOVELLA
349
isotopic signature is analogous to that of modern,
symbiotic species) (Kelly and others, 1998).
REPOSITORY.— Holotype (USNM 494824) and
paratype (USNM 494825) deposited at the Smithsonian
Museum of Natural History, Washington, D.C.
Morozovella apanthesma (Loeblich and Tappan,
1957)
DISCUSSION.— This taxon was discussed in the Atlas
of Paleocene Planktonic Foraminifera (Olsson and
others, 1999).
STRATIGRAPHIC RANGE.— Zone P3b to Zone E2.
In the Paleocene Atlas (Olsson and others, 1999) the
range of apanthesma was given as Zone P3b to P4c.
However, we have found this taxon in Zone E1 (=the
CIE interval) in the Bass River section of coastal New
Jersey and in several sections in Egypt and to extend to
near the top of Zone E2 (given as Zone P5c in papers in
Ouda and Aubry, 2003).
Morozovella aragonensis (Nuttall, 1930)
PLATE 11.3, FIGURES 1-16
(Pl. 11.3, Figs. 1-3: new SEMs of lectotype of
Globorotalia aragonensis Nuttall)
Globorotalia aragonensis Nuttall, 1930:288, pl. 24: figs. 6-
8, 10-11 (10-1l: lectotype USNM 59500 selected by Blow,
1979:990) [lower Eocene Aragon Fm., La Antigua, Rio
La Puerta, Tampico Embayment, Mexico].—Glaessner,
1937:10, pl. 1: figs. 5a-c [lower Eocene Lower
Foraminiferal Beds, Il’skaya, north west Caucasus].—
Subbotina, 1953:215, pl. 18: figs. 6a-c [Zone of conical
globorotaliids, Foraminiferal Beds, F1 unit, Nal’chik,
Khieu River section, North Caucasus]; ?7a-c [Zone of
compressed globorotaliids, lower White (subfish) Fm.,
Kolodets Koshak, Mangyshlak Peninsula, former soviet
union].—Bolli, 1957a:75, pl. 18, figs. 7-9 [G. aragonensis
Zone, upper Lizard Springs Fm., Trinidad].—Bolli,
1957b:167, pl. 38: figs. 1a-c [middle Eocene Hantkenina
aragonensis Zone, Navet Fm., Trinidad].—Mallory,
1959:252, pl. 35: figs. 2a-c [lower Eocene Lodo Fm.,
Ulatisian Stage, Devils Den, Fresno Co., California].—
Luterbacher, 1964:696-698, figs. 121a-c, 122a-c [G.
aragonensis Zone]; figs. 123a-c, 124a-c, 125a-126c [G.
bullbrooki Zone, Gubbio section, Apennines, Italy].—
Postuma, 1971:172, figs. on p. 173 [topotypes from La
Antigua, Rio La Puerta, Tampico region, Mexico].—
Luterbacher, 1975a:726, pl. 2: figs. 5a-c [Globorotalia
aragonensis Zone, DSDP Site 305, South Atlantic
Ocean].—Toumarkine, 1975:736 (not listed on p. 736 in
10:CC; compare Table 2, p. 737), pl. 2, figs. 9,10 [probably
Zone E9, but given as “G. lehneri to T. r ohri Zone”, DSDP
Site 305, South Atlantic Ocean].—Cifelli and Belford,
1977:102-103, pl. 1: figs. 7-9 [lectotype, CC NO. 64893
designated; lower Eocene Aragon Fm., La Antigua, Rio
La Puerta, Tampico Embayment, Mexico, referred to
Truncorotaloides (Morozovella)].—Toumarkine and
Luterbacher, 1985:112, text-figs. 16.4a-c [=Nuttall, 1930,
pl. 24, figs. 6-8 reillustrated; not holotype as stated; cf.
Cifelli and Belford, 1977:102-103 and Blow, 1979:990
who chose other illustrated specimen(s) as lectotype(s)];
16.5a-c [reillustration of Bolli, 1957a, pl. 18, figs. 7-9];
16.6a-c [Subbotina, 1953, pl. 18: figs. 6a-c reillustrated].
Globorotalia (Truncorotalia) aragonensis Nuttall.—Cushman
and Bermúdez, 1949:38, pl. 7: figs. 13-15 [Zone P9=E7,
Universidad Fm., Avenida de los Presidentes, Vedado,
Havana, Cuba].
Pseudogloborotalia aragonensis (Nuttall).—Bermúdez,
1961:1338-1340, pl. 16: figs. 5a-c [lower Eocene Zone
P9, Universidad Fm., Havana, Cuba].
Morozovella aragonensis aragonensis (Nuttall).—Fleisher,
1974:1029, pl. 14: fig. 11 [Zone P11, DSDP Site 219,
Indian Ocean].
Morozovella aragonensis (Nuttall) .—Berggren, 1977:244,
Chart No. 9, (various forms reillustrated from literature).—
Pearson, and others, 1993:124, pl. 2: figs. 1-3 [Zone P11,
DSDP Site 523, South Atlantic Ocean].—Warraich,
Plate 11.2, 1-3, Morozovella velascoensis (Cushman, 1925); 4-16, Morozovella allisonensis (Kelly, Bralower and Zachos, 1998)
Morozovella velascoensis 1-3, Zone E1, Bass River Borehole, New Jersey, ODP 174AX, 1169.1-.2 feet. Morozovella allisonensis 4-8, 12
(5-7, holotype, USNM 494824; 4, 8, 12, paratype, USNM 494825), Zone E1, ODP Hole 865C/12H/4, 10-12 cm, Allison Guyot, central
equatorial Pacific Ocean; 9-11, 13-16, Zone E1, Bass River Borehole, New Jersey, ODP 174AX, 1167.8-1171.2 feet. Scale bar: 1-16 = 100
µm.
BERGGREN AND PEARSON
350
Ogasawara and Nishi, 2000:293, fig. 17. 6,11,12 [Zone
P7, Dungan Fm., Rakhi Nala River, Sulaiman Range,
Pakistan].—Warraich and Ogasawara, 2001:37 [Zone P7,
Dungan Fm., Rakhi Nala River, Sulaiman Range,
Pakistan].Pearson and others, 2004:37, pl. 2, fig. 12
[middle Eocene, Zone P11, Tanzania Drilling Project Site
2, Kilwa Masoko, Tanzania].
Globorotalia (Morozovella) aragonensis Nuttall.—Blow,
1979:990, pl. 134: fig. 6 [Zone P8b, DSDP Hole 47.2,
Shatsky Rise, northwest Pacific Ocean]; pl. 141: figs. 1,2,
pl. 226, fig. 6 (detail of pl. 141, fig. 2) [Zone P8b, DSDP
Site 5, Atlantic Ocean] and figs. 4-9 [Zone P8b, DSDP
Site 5, Atlantic Ocean]; pl. 146, figs. 1, 2 [Zone P9, DSDP
Hole 47.2, Shatsky Rise, northwest Pacific Ocean]; pl.
147: figs. 5-7; pl. 152: figs. 1-5 [Zone P9, Kane 9-C piston
core, Endeavour Seamount, equatorial Atlantic Ocean];
pl. 164: figs. 6-9; pl. 227, figs. 1-3; pl. 228: figs. 4,5 [Zone
P10, Kane 9-C piston core, Endeavour Seamount,
equatorial Atlantic Ocean]; pl. 167: figs. 6-9; pl. 227: fig.
4; pl. 228, figs. 1-3; pl. 228: fig. 6 [Zone P11, Sample
RS24, Kilwa area, Tanzania].
Globorotalia marksi Martin, 1943:115, pl. 8: figs. 1a-c
[holotype, No. 7402, lower Eocene M. aragonensis Zone,
Lodo Fm., Lodo Gulch, Fresno County, California].
Globorotalia naussi Martin, 1943:116, pl. 8: figs. 5-6
[holotype No. 7403, Lodo Fm type section, Lodo Gulch,
Fresno Co., California].
DESCRIPTION.
Type of wall: Muricate, nonspinose, normal
perforate.
Test morphology: Test periphery nearly circular,
weakly lobulate, planoconvex; 5-7 chambers in last
whorl, triangular and inflated on umbilical side,
trapezoidal to lozenge shaped on spiral side as a function
of the strong curvature of intercameral sutures which
are raised or flush with test surface, muricate/beaded
and forming distinct, acute angle with periphery;
intercameral sutures staight to slightly sinuous and
moderately depressed on umbilical side; umbilicus
narrow, deep, and rimmed by rounded tips at
circumumbilical chamber confluence; blunt-tipped
muricae scattered over chambers of last whorl on
umbilical side; terminal chamber generally smooth;
strong/thick peripheral muricocarina; early chambers
strongly muricate obscuring early whorl(s); muricae on
early whorl(s) of test preclude accurate estimate of
number of chambers and whorls on test; probably about
12-15 in about 3 whorls; in edge view umbilico-convex;
spiral side flat or nearly so; primary aperture a low,
umbilical-extraumbilical arch extending to peripheral
margin.
Size: Lectotype dimension(s): not given by
Cifelli and Belford (1977); average diameter: 0.6 mm
(Nuttall, 1930, p. 288).
DISTINGUISHING FEATURES.— This taxon is
characterized by its tightly coiled, nearly circular test
(giving the appearance of a truncated cone), narrow
umbilicus, distinctly trapezoidal chambers on spiral side
and strongly developed muricae on the early chambers.
DISCUSSION. Nuttall (1930) did not designate a
holotype for this species when he described it from the
lower-middle Eocene Aragon Formation of Mexico. In
an examination of various species of Cenozoic
planktonic foraminiferal taxa in the collections of the
USNM, Cifelli and Belford (1977, p. 102, 103, pl. 1,
figs. 7-9) designated one (PP 64893) of 5 paralectotypes
in Nuttall’s original material (No. CC59499) as lectotype
and provided a detailed description of its morphologic
characters. The taxonomic characters of M. aragonensis
were also discussed by Blow (1979, p. 990-993) who
also attempted to designate a lectotype (USNM 59500)
for the taxon. However, there is some confusion
regarding Blow’s actions in this respect. There is also
considerable confusion over Blow’s action in its own
right because he did not adequately identify the
Plate 11.3 Morozovella aragonensis (Nuttall, 1930)
1-3 (lectotype, USNM 64893), lower Eocene, Aragon Fm., Vera Cruz, Tampico Embayment, Mexico; 4, Zone E8, ODP Hole 865C/8H/3,
110-112 cm, 8, 12, Zone E8, ODP Hole 865B/6H/4, 89-91 cm. Allison Guyot, equatorial Pacific Ocean; 5, Zone E9, TDP Site 13/16/1, 65-
75 cm, Mkazambo, near Pande, Tanzania; 6, Zone E9, TDP Site 3/17/2, 35-43 cm, Kilwa Masoko, Tanzania; 7 (Blow, 1979, pl. 164, fig.
6), Zone E8, KANE 9-Core 42, 15 cm, 9, 10, 15 (Blow, 1979, pl. 152, figs. 1, 3, 4), Zone E7, KANE 9-Core 42, 163 cm, Endeavor
Seamount, eastern North Atlantic Ocean; 11, Zone E8, TDP Site 2/18/1, 41-52 cm, 13 (reillustration of pl. 2, fig. 12, Pearson and others,
2004), Zone E8, TDP Site 2/17/2, 35-43 cm, 16, Zone E8, TDP Site 2/19/1, 10-20 cm, Kilwa, Tanzania; 14 (Blow, 1979, pl. 146, fig. 1),
Zone E7, DSDP 47/7/2, 65-67 cm, Shatsky Rise, northwest Pacific Ocean;. Scale bar: 1-16 bar = 100 µm.
CHAPTER 11 - MOROZOVELLA
351
PLATE 11.3 Morozovella aragonensis (Nuttall, 1930)
BERGGREN AND PEARSON
352
PLATE 11.4 Morozovella caucasica (Glaessner, 1937)
CHAPTER 11 - MOROZOVELLA
353
collection number of the desired specimen. Blow (1979,
p. 990) stated that Nuttall (1930) had “only selected two
cotypes which were deposited in the Cushman Collection
(numbers 59500 and 59499). The writer here selects the
specimen registered as Cushman Collection no. 59500
as lectotype of the taxon Globorotalia aragonensis
Nuttall 1930; this specimen is believed to be the one
figured by Nuttall in dorsal and ventral views, on his
plate 24, figs. 10 and 11…” However, we have been
able to verify (Brian Huber, personal communication
to WAB November 11/12, 2002) that there are currently
4 paralectotype specimen’s in Nuttall’s original CC59499
slide and 14 specimens in CC 59500! Since Cifelli and
Belford selected a lectotype from a slide with 5
specimens it must have come from CC 59499. The
mystery remains why Blow (1979, p. 990) stated that
Nuttall (1930) had deposited only two “cotypes “ in
the Cushman Collections. We suspect the answer may
lie somewhere in the following: 1) Blow (1979, p. 990)
may have chosen his lectotype without actually viewing
the slide(s) in question at the USNM; 2) Blow (1979, p.
990) selected what he believed to be a single specimen
illustrated by Nuttall (1930, pl. 24, figs. 10,11) to serve
as the lectotype of Globorotalia aragonensis Nuttall;
3) he would have considered that figures 6-8, pl. 24 in
Nuttall (1930) were three views of the second specimen
in the slide; 4) Blow (1979, p. 990) considered that the
“specimen” he had chosen as lectotype corresponds to
CC no. 59500, whereas the other “specimen” (which he
believed to correspond to pl. 24, figs. 6-8) corresponds
to CC no. 59499; 5) however, the five illustrations (CC
no. 59499) of Nuttall (1930, pl. 24, figs. 6-8, 10,1l) that
Blow (1979, p. 990) believed to represent two specimens
are, in actual fact, five specimens (see above and also
Cifelli and Belford, 1977: “The type material consists
of five cotypes that were figured in the original
description and another group of cotypes that were
unfigured” [CC no. 59500]); 6) It is thus obvious that
Blow (1979) and Cifelli and Belford (1977) chose their
respective lectotypes from the same sample (CC no.
59499) containing the five specimens deposited by
Nuttall in 1930. Blow (1979, p. 990), however, had
designated his lectotype based on two specimens; 7) it
is difficult to state/determine which of the five specimens
Cifelli and Belford (1977) designated as lectotype of
aragonensis, although a comparison of their illustration
(1977, pl. 1, figs. 7-9) with the five figured specimens
of Nuttall (1930, pl. 24, figs. 6-8, 10,1l) suggests it may
well be that figured by Nuttall (1930, pl. 24, fig. 1)
inasmuch as the illustration(s) of the spiral side of that
specimen shows a strong similarity to that of Cifelli and
Belford (1977, pl. 1, fig. 7). Blow had died in 1972 and
his monograph did not appear (1979) until two years
after the publication of the Cifelli and Belford paper
(1977). Thus, his designation of a lectotype, while
intriguing, is irrelevant to the taxonomic status of
aragonensis.
Blow (1979, p. 990) indicated that he considered
lensiformis to have been the direct ancestor of both crater
and aragonensis. We agree with this viewpoint. Blow
(1979, p. 1006, 1007) also considered marksi Martin and
naussi Martin as “ex interc. lensiformis and
aragonensis and as “primitive aragonensis”,
respectively, based on an examination of paratypes in
the USNM. Examination of the holotypes of these two
taxa (WAB) suggests that the morphologic differences
between these forms and “typical” aragonensis are of a
minor nature and that separate taxic distinction is not
warranted.
Morozovella aragonensis is one of the most
distinctive and widely cited morozovellids in Eocene
literature.
PHYLOGENETIC RELATIONSHIPS. This taxon
evolved from M. lensiformis and does not appear to have
left any descendants.
Plate 11.4 Morozovella caucasica (Glaessner, 1937)
1, 2, 5-7 (1, 2 and 5-7 same specimens), lower Eocene, northwestern North Atlantic Ocean; 3, 4 (same specimen), Zone E7, La Tuilerie de
Donzaque, France; 8, Zone E7, ODP Hole 865B/9H/1, 60-62 cm, 9, 11, Zone E8, ODP Hole 865C/8H/3, 110-112 cm, Allison Guyot,
equatorial Pacific Ocean; 10, 15 (Blow, 1979, pl. 152, figs. 8, 9), Zone E7, KANE 9-Core 42, 163 cm, 12, 14 (Blow, 1979, pl. 147, figs.
9, 10), Zone E7, KANE 9-Core 42, 200 cm, Endeavor Seamount, equatorial Atlantic Ocean; 13, 16, Zone E7, Tercis, Lesperon, Spain.
Scale bar: 1-16 = 100 µm.
BERGGREN AND PEARSON
354
STRATIGRAPHIC RANGE. Base of Zone E5 (by
definition) to top of Zone E9 (by definition).
GEOGRAPHIC DISTRIBUTION. Widely distrib-
uted in (sub)tropical-Tethyan regions; common in
Caribbean, Mediterranean-Pyrenees, North Caucasus,
Indo-Pacific, among others.
STABLE ISOTOPE PALEOBIOLOGY. Oxygen and
carbon isotopes indicate a shallow mixed-layer habitat
(Boersma and others, 1987; Pearson and others, 1993,
2001).
REPOSITORY. Lectotype (PP 64893) deposited at the
Smithsonian Museum of Natural History , Washington,
D.C. (see Cifelli and Belford, 1977).
Morozovella caucasica (Glaessner, 1937)
PLATE 11.4, FIGURES 1-16
Globorotalia aragonensis Nuttall var. caucasica Glaessner,
1937:31 (English):48 (Russian), pl. 1, figs. 6a-c [lower
Eocene upper beds of the lower part of the Lower
Foraminiferal (Koun) Fm., Il’skaya, north west Caucasus,
Former Soviet Union].
Truncorotalia caucasica (Glaessner).—Reiss, 1957:239-241
(comments on taxonomy).—Bykova, Vasilenko,
Voloshinova, Myatliuk and Subbotina, 1959:302-303, text-
figs. 691A-C (reillustration of Subbotina, 1953, pl. 19,
figs. 2a-c) [Zone of conical globorotaliids, Kuban River
section, Foraminiferal Beds, Green Fm., North Caucasus,
Former Soviet Union].
Truncorotalia (Truncorotalia) cf. caucasica Glaessner.—von
Hillebrandt, 1962:136-137, pl. 13: fig. 5 [Zone G,
Reichenhall-Salzburg Basin, Germany].
Globorotalia caucasica Glaessner—Luterbacher, 1964:684,
fig. 97 [Zone of conical globorotaliids, Khieu River, North
Caucasus, Former Soviet Union].—Stainforth and others,
1975:175-176, text-fig. 41: 1-2 (1: from Subbotina,
1953:19, figs. 1a-c; fig. 2 ascribed incorrectly to
Subbotina, 1953), pl. 19, fig. 2 (it is actually fig. 3 and
indicated as transitional to aragonensis); 41: 3-6 [lower
Eocene G. pentacamerata Zone, northern Caucasus,
former Soviet Union].—Luterbacher, 1975b:66, pl. 4, figs.
28-30 [Globorotalia pentacamerata Zone, Possagno
section, Italy].
Globorotalia (Morozovella) crater caucasica Glaessner.—
Jenkins, 1971:103, text-figs. 189-191 [G. (M.) crater Zone,
Mangaorapan sample N141/921, Waipawa Standard
Section, North Island, New Zealand].
Morozovella aragonensis caucasica (Glaessner).—Fleisher,
1974:1029, pl. 14: fig. 2 [Zone P8, DSDP Site 220, Indian
Ocean].
Globorotalia (Morozovella) caucasica Glaessner.—Blow,
1979:993, pl. 146: figs. 3 and 4 [Zone P9, DSDP Hole
47.2, Shatsky Rise, northwest Pacific Ocean]; pl. 147: figs.
9, 10; pl. 226: fig. 1; pl. 152: figs. 6-9; pl. 226: fig. 2; pl.
226: figs. 3,4 [Zone P9, Kane 9-C piston core, North
Atlantic Ocean].
Morozovella caucasica (Glaessner).—Toumarkine and
Luterbacher, 1985:114, figs. 16.2-3 [2: from Subbotina,
1953, pl. 19: fig. 1; 3: holotype reillustrated].
Globorotalia velascoensis (Cushman).—Subbotina,
1953:216-219, pl. 19, figs. 1-2 [Zone of conical
globorotaliids, Foraminiferal beds, Green Fm., Kuban
River section, North Caucasus, Former Soviet Union]; fig.
3 [Zone of conical globorotaliids, Kutais Horizon [F3],
Foraminiferal Beds, Gubs River section, Barakaev region,
North Caucasus, Former Soviet Union]. [Not Cushman,
1925.]
DESCRIPTION.
Type of wall: Muricate, nonspinose, normal
perforate.
Test morphology: Test subcircular, moderately
lobulate peripheral outline, planoconvex; chambers
triangular on umbilical side, trapezoidal to subquadrate
on spiral side as a function of degree of curvature of
Plate 11.5 Morozovella crater (Hornibrook, 1958)
1, 2, 13 (Blow, 1979, pl. 138, figs.. 6, 7, 8), Zone E6, DSDP Hole 20C/5/5, 72-74 cm, central South Atlantic Ocean; 3, 4, 6, 8 (Blow, 1979,
pl. 134, figs. 1, 2, 4, 5), Zone E6, DSDP Site 47:2/8/1, 77-79 cm; 5, 12 (Blow, 1979, pl. 164, figs. 3, 4), Zone E8, KANE 9-Core 42, 15
cm; 7 (Blow, 1979, pl. 147, fig. 8), Zone E7, KANE 9-Core 42, 200 cm, Endeavor Seamount, equatorial Atlantic Ocean; 9-11 (same
specimen), lower Eocene, DSDP Site 384, northwestern North Atlantic Ocean; 14 (Blow, 1979, pl. 146, fig. 5), Zone E7, DSDP Site 47:2/
7/2, 65-67 cm, Shatsky Rise, northwest Pacific Ocean; 15, 16, Zone E8, DSDP Site 94/30/2, 37-38 cm, Gulf of Mexico. Scale bar: 1-16 =
100 µm.
CHAPTER 11 - MOROZOVELLA
355
PLATE 11.5 Morozovella crater (Finlay, 1939)
BERGGREN AND PEARSON
356
PLATE 11.6 Morozovella edgari (Premoli Silva & Bolli, 1973)
CHAPTER 11 - MOROZOVELLA
357
intercameral sutures; muricae strongly developed on
circumumbilical collar; primary aperture a low
umbilical-extraumbilical slit extending to peripheral
margin; sutures depressed, straight, radial; umbilicus
large, deep surrounded by steeply plunging
circumumbilical chamber walls and surmounted by
thickly ornamented circumumbilical collar of fused
muricae which rim the everted margins of the chambers;
in spiral view 15-18 chambers arranged in approximately
3 whorls; early chambers slightly raised above test
surface; gradual increase in chamber size throughout;
muricate sutures strongly curved, flush with/slightly
elevated above test surface; in edge view umbilico-
convex; spiral side flat or nearly so; strongly muricate/
beaded keel.
Size: Holotype dimension(s): not given; range
given: 0.4-0.6 mm (diameter).
DISTINGUISHING FEATURES. This taxon is
characterized by having a strongly planoconical test with
6-8 (less commonly 5 at one extreme and 9-10 at the
other) chambers in the final whorl, a large open and deep
umbilicus whose everted circumumbilical chamber
margins are strongly ornamented by fused muricae. It is
strongly homeomorphic with M. velascoensis from the
late Paleocene-early Eocene (Zone P3b-E2), but M.
velascoensis s.s. may be distinguished by its (generally)
somewhat larger size, higher conical angle and more
pronounced circumumbilical ornament of everted
chamber shoulders (tips).
DISCUSSION. There exists a variety of opinions on
the phylogenetic affinities of this taxon which may be
summarized as follows:
1. Although Glaessner (1937) clearly
distinguished between his new taxon caucasica (early
Eocene) and velascoensis Cushman (late Paleocene), the
two forms were frequently confused in the literature (see
Subbotina, 1953 and El Naggar,1966, for example) as
well as aragonensis and caucasica (identified as
velascoensis; Subbotina, 1947; see Subbotina, 1953, p.
216). It was Reiss (1957; see also Blow, 1979) who
provided clear differentiation of the two morphotypes
on the basis of morphology and stratigraphic distribution.
Indeed, one of us (WAB) pointed out this differentiation
to Subbotina during a visit to VNIGRI (St. Petersburg)
in 1958 and by the late 1950s Subbotina had incorporated
the distinction between the two as shown by the correct
identification of caucasica in the Soviet Treatise on
Paleontology (Bykova and others, 1959).
2. Hillebrandt (1962) included crater Finlay and
formosa Bolli in the synonomy of caucasica and
considered it to have evolved from G. (T.) aequa
simulatilis (see Berggren and Norris, 1997, p. 78 for a
discussion of the probable/possible taxonomic affinities
of the late Paleocene taxon simulatilis).
3. The holotype of Globorotalia crater Finlay
is a 5-chambered morphotype similar in several respects
to caucasica. Hornibrook (1958) drew attention to the
“wide umbilicus surrounded by strongly muricate (not
true spines) distal ends of chambers” as characteristic
of crater. Jenkins (1971) considered M. caucasica a
subspecies of crater and to differ only in the number of
chambers ( 6-8 vs. 5) from the latter. Like Hillebrandt
(1962), Jenkins (1971) considered formosa a junior
synonym of crater.
4. Stainforth and others (1975) considered that
crater is possibly a “highly ornamented member of the
Globorotalia formosa formosa plexus and G. caucasica
is a homeomorph originating from Globorotalia
aragonensis with which it is linked by transitional
forms”. In support of this view Stainforth and others
(1975) observed that in New Zealand G. crater is used
to delineate a zone which extends over/correlates with
the G. formosa formosa, G. aragonensis and G.
Plate 11.6 Morozovella edgari (Premoli Silva and Bolli, 1973)
1-7 (3, holotype, Premoli Silva and Bolli, 1973, pl. 8, fig. 1; 1, 5-7, paratypes, pl. 8, figs. 2, 4, 7, 9; figs.. 2, 4, paratypes, pl. 7, figs. 10, 12),
Zone E3, DSDP Site 152/3/2, 136-138 cm; 8-10, Zone E4, ODP Hole 865B/11H/3, 57-59 cm; 11, (Kelly and others, 2001, fig. 3C), Zone
E3, ODP Hole 865B, 95.70 mbsf, Allison Guyot, equatorial Pacific Ocean; 12, 16, Zone E2, TDP Hole 7A/65/2, 71-86 cm, Kilwa,
Tanzania; 13-15 (Kelly and others, 2001, fig. 2E), Zone E4, DSDP Site 152/3/5, 21-23 cm, Caribbean Sea. Scale bar: 1-16 = 100 µm.
BERGGREN AND PEARSON
358
pentacamerata Zones, that is, a zone which extends
below the base of the typical 6-8 chambered caucasica.
5. Blow (1979) extended these views in
separating crater (5-chambered) and caucasica (6-8
chambered; see also Fleisher, 1974, pl. 14, fig. 2 who
illustrated a morphotype with apparently 9 or 10
chambers from Zone P11 of the Indian Ocean) on the
basis of chamber number and the narrower umbilicus of
crater. He asserted that crater lies much closer to the
caucasica end of the lensiformis-crater-caucasica
lineage than it does to the ancestral lensiformis
morphotype, a view with which we can readily agree.
Thus, Blow (1979) considered crater as a form
transitional between lensiformis and caucasica with
typical 5-chambered morphotypes extending into
horizons as high as Zone P10 (middle Eocene).
Morozovella twisselmanni (Mallory) was considered a
junior synonym of crater as well. However, Blow (1979)
also illustrated several morphotypes which he attributed
to crater but which also bear a close resemblance to
formosa Bolli: 5-6 chambers, narrow umbilicus, absence
of circumumbilical muricate ornament ( see for instance
forms which he considered typical of crater by
comparison with paratypes of crater; Blow, 1979, pl.
138, figs. 4-8).
6. Several authors have considered a
phylogenetic relationship between aragonensis and
caucasica more likely (Fleisher, 1974; Berggren 1977;
Toumarkine and Luterbacher, 1985).
Our own studies suggest that caucasica is,
indeed, more closely related to crater and that it
represents the end member of the subbotinae-
lensiformis-crater-caucasica lineage, whereas
aragonensis is the end member of a divergent M.
subbotinae-lensiformis-aragonensis lineage.
PHYLOGENETIC RELATIONSHIPS.— This taxon is
the end member of the M. subbotinae-lensiformis-crater-
caucasica lineage and does not appear to have left any
descendants. It is strongly homeomorphic with, but
unrelated in terms of lineal descent to, M. velascoensis.
STRATIGRAPHIC RANGE.— Zone E6 to Zone E8.
GEOGRAPHIC DISTRIBUTION.— Widely distributed
in (sub)tropical-Tethyan regions; particularly common
in the Aquitaine Basin (France), the Indo-Pacific region,
among others.
STABLE ISOTOPE PALEOBIOLOGY.— No data
available.
REPOSITORY.— Holotype may have been deposited
in the collections of the Paleontology Department,
University of Moscow; not seen during visits in 1962,
1973 and 1988 (WAB).
Morozovella crater (Hornibrook, 1958)
PLATE 11.5, FIGURES 1-16
Globorotalia crater Finlay, 1939a:125, (no figures; nomen
nudum).—Hornibrook, 1958:33, pl. 1: figs. 3-5 [holotype
figured and redescribed from lower Eocene, sample F
5570, chalk marls below Amuri Strone, Hurunui River,
Heretaungan Stage, New Zealand].
Globorotalia (Morozovella) crater Finlay.—Jenkins,
1971:103, pl. 8: figs. 192-194 (refigured holotype), figs.
195-197 [lower Eocene, middle Waipara River section,
Mangaorapan Stage, New Zealand].—Blow, 1979:996. pl.
134: figs. 2-5, 9 and 10; pl: 135, figs. 1,2; pl: 224, figs.
5,6 [detail of pl. 135: fig. 2] [Zone P8b, DSDP Hole 47.2,
Shatsky Rise, northwest Pacific Ocean]; pl. 138: figs. 4-
8; pl. 225: fig. 1 [ Zone P 8b, DSDP Hole 20C, South
Atlantic Ocean]; pl. 141: fig. 3; pl. 146: fig. 5 [Zone P9,
DSDP Hole 47.2, Shatsky Rise, northwest Pacific Ocean];
pl. 154: figs. 2-4; pl. 225: figs. 3-6; pl. 164: figs. 3-5 [Zone
P10; Kane 9-C piston core, Endeavour Seamount,
Plate 11.7 Morozovella formosa (Bolli, 1957)
1-3 (holotype, USNM P5056), Zone E5, upper Lizard Springs Fm., Trinidad; 4, lower Eocene, Khieu River, northern Caucasus; 5-7, Zone
E6, ODP Hole 865B/10/1, 62-64 cm, 7, 8, 13, Zone E4, ODP Hole 865B/11H/3, 57-59 cm, 9-12, 14-16 (9-11; 12, 16; 14, 15 same
specimens), Zone E4 ODP Hole 865B/11H/3, 57-59 cm, Allison Guyot, equatorial Pacific Ocean. Scale bar: 1-16 = 100 µm.
CHAPTER 11 - MOROZOVELLA
359
PLATE 11.7 Morozovella formosa (Bolli, 1957)
BERGGREN AND PEARSON
360
PLATE 11.8 Morozovella gracilis (Bolli, 1957)
CHAPTER 11 - MOROZOVELLA
361
equatorial Atlantic Ocean]; pl. 226: fig. 5 [Zone P8b Hole
20C, South Atlantic Ocean].
Morozovella crater (Finlay). Pearson and others, 2004:37,
pl. 2, figs. 13, 14 [middle Eocene, Zone P11, Tanzania
Drilling Project Site 2, Kilwa Masoko, Tanzania].
Globorotalia aragonensis Nuttall var. twisselmanni.—
Mallory, 1959:252, pl. 23: figs. 1a-c [lower Eocene,
Penutian Stage, lower Lodo Fm., Media Agua Creek,
California].
Globorotalia (Morozovella) gorrondatxensis Orue-Etxebarria,
1985:471, pl. 2: figs. 1-10 [lower middle Eocene
Truncorotaloides praetopilensis Zone, Getxo, westernmost
part of Gorrondatxe Beach to Azkorri Beach, Vizkaya
Province, Spain].
Not Globorotalia crater Finlay, 1939b: pl. 29: figs. 157, 162,
163 [upper Paleocene, mid-Waira, New Zealand].
DESCRIPTION.
Type of wall: Muricate, nonspinose, normal
perforate.
Test morphology: Plano-convex test with lobulate
outline, ornamented with well developed muricocarina;
41/2-5 essentially equidimensional chambers in last
whorl; umbilical intercameral sutures weakly curved to
radial; sutures on spiral side curved, limbate, heavily
ornamented (beaded); umbilicus deep and relatively
wide, rimmed by everted, thickened circumumbilical rim
of elevated chamber shoulders (tips); in edge view
distinctly plano-convex; aperture a low, interiomarginal,
umbilical-extraumbilical slit with thin lip.
Size: Maximum diameter of holotype: 0.53 mm
(Jenkins, 1971, p. 104).
DISTINGUISHING FEATURES.— 41/2 to 5 essentially
equidimensional chambers in last whorl, thickened
circumumbilical rim of elevated chamber shoulders,
strongly limbate sutures on spiral side.
DISCUSSION. In erecting the new species
Globorotalia crater Finlay (1939a) gave a brief
description but no formal illustration(s). Thus, G. crater
was “nomen nudum”. Specimens figured by Finlay
(1939b, pl. 29: figs. 157, 162, 163) are from older strata
than the type level and are of a different species
(Hornibrook, 1958). The first valid reference to M. crater
is by Hornibrook (1958, p. 33, pl. 1, figs. 3-5) in which
a holotype specimen is illustrated and a more thorough
description given. It would appear, then, that Hornibrook
(1958) is to be credited with authorship of the species
crater rather than Finlay (1939 a, b; see also Berggren,
1977, p. 245, 246). Jenkins (1971, p. 103) and Blow
(1979, p. 996) have presented extensive discussion of
this taxon and suggested that it is either a junior synonym
(Jenkins, 1971) or a pene-contemporaneous
homeomorph (Blow, 1979) of M. formosa. We have
found in the course of our work that crater exhibits a
general test shape closer to that of aragonensis
(subcircular, weakly lobulate periphery) than formosa
(moderately to strongly lobulate periphery and early
vorticiform spiral intercameral sutures).
PHYLOGENETIC RELATIONSHIPS. This form
evolved from M. lensiformis and evolved into M.
caucasica.
STRATIGRAPHIC RANGE. Zone E4 to Zone E9.
GEOGRAPHIC DISTRIBUTION. Widely distributed
in (sub)tropical areas of Atlantic, Mediterranean/
Tethyan, Pacific Oceans and austral regions.
STABLE ISOTOPE PALEOBIOLOGY. No data
available.
REPOSITORY. Holotype (New Zealand Geological
Survey Register No. TF 1077) refigured and described
by Hornibrook (1958), deposited in New Zealand
Geological Survey, Lower Hutt, N.Z.
Plate 11.8 Morozovella gracilis (Bolli, 1957)
1-3 (holotype, USNM P5055), Zone E4, upper Lizard Springs Fm., Trinidad; 4, 8, 12-16, Zone E4, ODP Hole 865B/11H/3, 57-59 cm,
Allison Guyot, equatorial Pacific Ocean; 5-7, 9-11 (same specimens), uppermost Paleocene, DSDP Site 384, northwestern North Atlantic
Ocean. Scale bar: 1-16 = 100 µm.
BERGGREN AND PEARSON
362
Morozovella edgari (Premoli Silva and Bolli, 1973)
PLATE 11.6, FIGURES 1-16
(Pl. 11.6, Figs. 1-7: reillustrations of holotype and
paratypes of Globorotalia edgari Premoli Silva and
Bolli)
Globorotalia edgari Premoli Silva and Bolli 1973:526, pl. 7:
figs. 10-12; pl. 8: figs. 1 (holotype)-12 [Globorotalia
edgari Zone, DSDP Site152, Nicaragua Rise, Caribbean
Sea].
Morozovella edgari (Premoli Silva and Bolli).—Toumarkine
and Luterbacher, 1985:114, text-figs. 15.6a (holotype) and
6b, c (paratypes) reillustrated.—Kelly and others,
2001:509, text-fig. 1D-F and text-fig. 3C [lower Eocene,
ODP Hole 865B, Allison Guyot, Mid-Pacific Mountains,
Pacific Ocean]; text-fig. 2E [Zone P6a, DSDP Site 152,
Caribbean Sea].
Globorotalia (Morozovella) finchi Blow, 1979:999, pl. 99:
figs. 6-11 (holotype) [Zone P5, DSDP Hole 47.2, Shatsky
Rise, Pacific Ocean].
DESCRIPTION.
Type of wall: Muricate, nonspinose, normal
perforate.
Test morphology: Low to moderately conical
trochospiral; weakly lobulate peripheral outline, 5-6
gradually enlarging , cuneiform-shaped chambers in last
whorl, intercameral sutures on umbilical side radial,
straight and weakly incised; on spiral side chambers
trapezoidal in shape, separated by distinctly curved
sutures which are flush with the test; early part/chambers
of the test raised (i.e., the test exhibits an unequal
biconvexity); umbilicus narrow, deep; distinct
muricocarina extends to final chamber; aperture a low
umbilical-extraumbilical arch extending nearly to the
periphery; surface distinctly muricate with concentration
of pustules decreasing towards terminal chambers.
Size: Dimensions of holotype: maximum
diameter: 0.21 mm.
DISTINGUISHING FEATURES.— This taxon is
characterized by its relatively small size, narrow and
deep umbilicus, and weakly developed peripheral
muricocarina. It bears a close resemblance to juvenile
individuals of M. velascoensis from which it is believed
to have evolved (Kelly and others, 2001).
DISCUSSION.— This minute morozovellid (test
diameter ranges from 0.2-0.25 mm) is associated with,
and ranges beyond, terminal members of the
Morozovella velascoensis group. Kelly and others (2001,
p. 507) have suggested that edgari descended from
velascoensis by a process called “terminal progenesis”
in which the diminutive descendant (edgari) with an
adult morphology resembles the juvenile stages of the
ancestral form velascoensis.
Morozovella finchi Blow is placed in the
synonomy of this taxon. It shares most morphologic
features in common with edgari except that the holotype
is somewhat larger (0.32 mm) in diameter. It was
described from Zone P5 (of Blow, 1979) and said to
range to Zone P7 (=E4 of this paper), a range comparable
to that ascribed to edgari here. Blow (1979, p. 999-1000)
was of the opinion that finchi descended from
Acarinina” trichotrocha and even included a paratype
from the Hornerstown Formation (Zone P4) of New
Jersey (erroneously ascribed to Zone P5) in his new taxon
finchi, but our examination of this form at the USNM
suggests that this specimen is not ascribable to finchi
(=edgari).
PHYLOGENETIC RELATIONSHIPS.— This taxon
evolved from Morozovella velascoensis (by means of a
gradual decrease in test size and truncation of the
Plate 11.9 Morozovella lensiformis (Subbotina, 1953)
Morozovella lensiformis 1-4 (Blow, 1979, pl. 126, figs. 5-8), Zone E6, DSDP Hole 47-2/8/2, 71-73 cm, Shatsky Rise, northwest Pacific
Ocean; 5-7, lower Eocene, Khieu River, northern Caucasus; 8-10, Zone E5, sample 914, bed 11, Khieu River, northern Caucasus; 11, 12,
16 (same specimen), lower Eocene, DSDP Site 39/2/CC, northeastern Pacific Ocean; 13-15, lower Eocene, DSDP Site 213, Indian Ocean.
Scale bar: 1-16 = 100 µm.
CHAPTER 11 - MOROZOVELLA
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PLATE 11.9 Morozovella lensiformis (Subbotina, 1953)
BERGGREN AND PEARSON
364
PLATE 11.10 Morozovella marginodentata (Subbotina, 1953)
CHAPTER 11 - MOROZOVELLA
365
ancestral ontogenetic sequence; Kelly and others, 2001)
but does not appear to have left any descendants.
STRATIGRAPHIC RANGE.— Zone E2 (uppermost
part) to Zone E3.
GEOGRAPHIC DISTRIBUTION.— Probably widely
distributed in (sub)tropical regions (Caribbean, Atlantic
Ocean and Pacific Ocean) but records are sparse owing
to this taxon having been overlooked heretofore. We have
found this form relatively commonly distributed in lower
Eocene strata (Esna Shales) of Egypt.
STABLE ISOTOPE PALEOBIOLOGY.— Shallow-
intermediate depth-habitat (Kelly and others, 2001).
REPOSITORY. Holotype (C-29482) and paratypes
(C- 29483 to C-29486) deposited at Natural History
Museum of Basel (Switzerland).
Morozovella formosa (Bolli, 1957b)
PLATE 11.7, FIGURES 1-16
(Pl. 11.7, Figs. 1-3: new SEMs of holotype of
Globorotalia formosa formosa Bolli)
?Globorotalia velascoensis (Cushman).Cushman and Renz,
1946:47, pl. 8: figs. 13-14 [upper Lizard Springs Fm.,
Trinidad; see Renz, 1951 for corrected figure explanation].
[Not Cushman, 1925.]
Globorotalia formosa formosa Bolli 1957b:76, pl. 18: figs.
1-3 [lower Eocene G. formosa formosa Zone, upper Lizard
Springs Fm., Trinidad].—Luterbacher, 1964:694-696,
text-figs. 118a-c [G. formosa formosa Zone, upper Lizard
Springs Fm., Trinidad]; 119a-c [G. formosa formosa/G.
subbotinae Zone, Gubbio section, Central Apennines,
Italy]; 120a-122c [G. aragonensis Zone, Gubbio section,
Italy].—Samanta, 1970:624, pl. 97: figs. 15, 16 [upper
Marlstone of Pondicherry Fm., southeastern India].—
Stainforth and others, 1975:184, text-fig. 48.1-5 [G.
formosa formosa Zone, Lodo Fm., California]; 48.6
[reillustration of holotype from Bolli, 1957b, pl. 18, figs.
1-3].—Luterbacher, 1975b:65, pl. 5: figs. 16-18 [G.
formosa formosa Zone, Possagno section, Italy].—
Toumarkine and Luterbacher, 1985:112, Fig. 15.13
[holotype reillustrated].
Pseuodogloborotalia formosa (Bolli).—Bermúdez,
1961:1343, 1344.
Globorotalia formosa Bolli.—Postuma, 1971:190, figures on
p. 191 [lower Eocene, Cuba].
Morozovella formosa (Bolli).—Berggren, 1971, pl. 5: figs.
15, 16 [Zone P6b, DSDP Hole 20C, South Atlantic Ocean].
Globorotalia (Morozovella) formosa Bolli.—Blow,
1979:1000, pl. 127: figs. 3 and 4 [Zone P8a, DSDP Hole
47.2, Shatsky Rise, northwest Pacific Ocean]; pl. 134:
figs. 7, 8, pl. 224: fig. 4 [Zone P8b, DSDP Hole 47.2,
Shatsky Rise, northwest Pacific Ocean]; pl. 138: figs. 9,
10; pl. 224: fig. 3 [Zone P8b, DSDP Hole 20C, South
Atlantic Ocean].
Morozovella formosa formosa (Bolli).—Warraich and others,
2000:293, fig. 17. 21-23 [Zone P7, Dungan Fm., Rakhi
Nala section, Sulaiman Range, Pakistan].—Warraich and
Ogasawara, 2001, 39, figs. 9. 13-15 [Zone P7, Dungan
Fm., Rakhi Nala section, Sulaiman Range, Pakistan].
DESCRIPTION.
Type of wall: Muricate, nonspinose, normal
perforate.
Test morphology: Test subcircular, moderately
lobulate peripheral outline, planoconvex; chambers
triangular, inflated and subangular on umbilical side;
trapezoidal with curved margins on spiral side; 15-18
chambers arranged in approximately 3 whorls on spiral
side and generally 6-61/2 chambers (rarely 7-8) in final
whorl on umbilical side with muricae scattered on early
chambers of last whorl; sutures moderately depressed,
weakly curved (early chambers) to straight, radial (later
chambers) on last whorl; umbilicus open, moderately
wide, deep; primary aperture a low umbilical-
Plate 11.10 Morozovella marginodentata (Subbotina, 1953)
1-3 (same specimen), Zone E5, sample 2911, 1 m above base of Cherkessk Fm., Khieu River, northern Caucasus; 4, 8, 12, lower Eocene,
Khieu River, northern Caucasus; 5-7 (same specimen), Zone E5, DSDP Site 98/10/1, 100-112 cm, Blake Plateau, western North Atlantic
Ocean; 9-11 (same specimen), Zone E4/5, DODO Piston Core 86P, 130-140 cm, Central Pacific Ocean; 13-16, Zone E4, ODP Hole 865B/
11H/3, 57-59 cm, Allison Guyot, equatorial Pacific Ocean. Scale bar: 1-16 = 100 µm.
BERGGREN AND PEARSON
366
extraumbilical arch extending to peripheral margin;
gradual increase in chamber size throughout although
last 2-3 chambers exhibit varying size; muricate sutures
strongly curved, flush with/slightly elevated above test
surface; umbilico-convex in edge view; spiral side flat
or nearly so; strongly muricate keel.
Size: Dimensions of holotype: maximum
diameter: 0.65 mm.
DISTINGUISHING FEATURES.— This taxon is
characterized by its relatively large, robust test, 6-7
(rarely 8) chambers and strongly muricate keel. It is
distinguished from its antecedent M. gracilis by its larger,
more robust test, larger number of chambers in the last
whorl and wider umbilicus, and from its partially
contemporaneous homeomorph, M. crater, by its more
lobulate periphery and larger number of chambers in the
final whorl and less ornamented test.
DISCUSSION.— Blow (1979, p. 1000, 1001) drew
attention to the fact that formosa and the crater-
caucasica complex, while seemingly morphologically
distinct when treated as isolated individuals, in fact
require close scrutiny in order to retain the
biostratigraphic utility of both. He further pointed out
the gross homeomorphy between formosa and some
morphotypes of the older velascoensis. Finally he
observed that some of the phylogenetically advanced
morphotypes of the velascoensis and the lensiformis-
crater-caucasica “gens” exhibit features in common with
formosa; a sort of “threefold homeomorphy” as he
dubbed it. Distinction among the different morphotypes
was made on the following basis:
1. formosa is flat to slightly convex in dorsal
(spiral) aspect with chambers somewhat longer (anterio-
posteriorly) tangentially than broad radially.
2. formosa has dorsal /spiral intercameral sutures
nearly radial proximally but recurved (but not sharply
retorse) near periphery.
3. velascoensis has chambers in dorsal/spiral
aspect nearly equidimensional and spiral intercameral
sutures exhibit retorse distal part.
4. In axial-apertural view anguloconical angle
in formosa is markedly acute (~ 45o), whereas in
velascoensis it is nearly a right-angle (~ 80-85o); the
umbilico-peripheral part of chamber is often somewhat
concave in the area between the peripheral test
muricocarina and the “horn-like”, adumbilically pointed
umbilical shoulders.
5. Whereas the anguloconical angle in the
crater-caucasica group is comparable to that in formosa,
the earlier chambers have a more rounded appearance
in axial/edge view.
6. The umbilicus in formosa is wide and the
angle made by the junction of the chamber walls interior
and exterior to the umbilicus is on the order of ~ 40-45o;
in crater this angle is ~ 60-65o in the case of the earlier
chambers and the umbilical shoulders are more gently
rounded, only the last 2-3 chambers exhibiting a strongly
reflexed angle between interior and exterior chamber
walls.
7. The main distinguishing feature separating
formosa from the other forms above is the nature of the
early vorticiform spiral intercameral sutures together
with the acute conical angle and the distinctly flat spiral
side.
PHYLOGENETIC RELATIONSHIPS.— This taxon
evolved from M. gracilis and does not appear to have
left any descendants.
STRATIGRAPHIC RANGE.— Base of Zone E4 (by
definition) to top of Zone E6 (by definition).
GEOGRAPHIC DISTRIBUTION.— Widely distributed
in (sub)tropical regions (Caribbean Sea, Atlantic, Pacific,
Tethyan, Indian and Austral Oceans).
STABLE ISOTOPE PALEOBIOLOGY. No data
available.
REPOSITORY. Holotype (USNM P5056) deposited
at the Smithsonian Museum of Natural History,
Washington, D.C.
Morozovella gracilis (Bolli, 1957b)
PLATE 11.8, FIGURES 1-16
(Pl. 11.8, Figs. 1-3: new SEMs of holotype of
Globorotalia formosa gracilis Bolli).
DISCUSSION.— This taxon was discussed in the Atlas
of Paleocene Planktonic Foraminifera (Olsson and
others, 1999).
STRATIGRAPHIC RANGE.— Zone P5 to Zone E5.
CHAPTER 11 - MOROZOVELLA
367
Morozovella lensiformis (Subbotina, 1953)
PLATE 11.9, FIGURES 1-16
Globorotalia lensiformis Subbotina, 1953:214, pl. 18: figs.
4a-c (holotype) and 5a-c [lower Eocene Zone of conical
globorotaliids, Foraminiferal Beds, Unit F1, Nal’chik,
Khieu River section, North Caucasus].—Luterbacher,
1964:673, text-figs. 74a-c [topotype from Zone of conical
globorotaliids, Khieu River section, northern Caucasus,
determined by N.N. Subbotina, fide Luterbacher].—
Samuel, 1972:192, 193, pl. 49: figs. 3-4c [middle Eocene,
Borehole Sr-1, Bakony Mountains, Hungary]; pl. 51: figs.
6a-b; 7a-b [Borehole Ot-69, Bakony Mountains,
Hungary].—Stainforth and others, 1975:200, text-figs. 1a-
2c (reillustrated from Subbotina, 1953); 3-7 [lower Eocene
G. formosa formosa Zone, Lodo Fm., California].
Globorotalia (Truncorotalia) lensiformis Subbotina.—von
Hillebrandt, 1962:136, pl. 13, figs. 12a-13c [Zone G,
Reichenhall-Salzburg Basin, Austria/Germany].
Morozovella lensiformis (Subbotina).—Berggren, 1971, pl.
5: figs. 18-20 [Zone P6b, DSDP Hole 20C, South Atlantic
Ocean].—Snyder and Waters, 1985:460, pl. 9: figs. 1, 3
[Zone P6b, DSDP Hole 548A, North east Atlantic
Ocean].—Warraich and others, 2000:293, fig. 17. 7-9
[Zone P7, Dungan Fm., Rakhi Nala section, Sulaiman
Range, Pakistan].—Warraich and Ogasawara, 2001:40,
figs. 10. 1-3 [Zone P7, Dungan Fm., Rakhi Nala section,
Sulaiman Range, Pakistan].
Globorotalia (Morozovella) lensiformis Subbotina.—Blow,
1979:1003-1005, pl. 125: figs. 6-9; pl. 126: figs. 1-3; pl.
128: figs. 1-9; pl. 129: figs. 1-3 [Zone P8a, DSDP Hole
47.2, Shatsky Rise, northwest Pacific Ocean]; pl. 134: fig.
7 and pl. 134: fig. 1 [Zone P8b, DSDP Hole 47.2, Shatsky
Rise, northwest Pacific Ocean]; pl. 251: fig. 5 [lower
Eocene, Region de Belu, France].
Globorotalia nartanensis Shutskaya, 1956:96-98, pl. 4: figs.
2a-c [G. subbotinae Zone, Cherkessk Horizon, Nal’chik,
central northern Caucasus, former Soviet Union].
Globorotalia californica Smith, 1957:190, pl. 28: figs. 22a-
23c [Vine Hill Sandstone, Selby, Contra Costa County,
California].—Mallory, 1959:253, pl. 38: figs. 4a-c [lower
Bulitian Stage, Lower Lodo Fm., Media Agua Creek,
California]. [The name Globorotalia californica Smith
1957 is a junior homonym of Globorotalia californica
Cushman and Todd 1948, which may be a
praeglobotruncanid from California; see Berggren, 1977,
p. 243.]
Globorotalia dolabrata Jenkins, 1965:1113, pl. 10: figs. 104-
112 (holotype) [lower Eocene M. crater Zone, upper part
of Waipawan Stage, Middle Waipara River section, North
Island, New Zealand].
Globorotalia (Morozovella) dolabrata Jenkins.—Jenkins,
1971:104, pl. 104: figs. 233-235 (holotype refigured); figs.
236 -238, 239-241 (paratypes) [lower Eocene M. crater
Zone, upper part of Waipawan Stage, Middle Waipara
River section, North Island, New Zealand].
Globorotalia (Morozovella) aequa dolabrata Jenkins.Blow,
1979:981, pl.125: figs. 3-5 and pl. 127: fig. 8 [Zone P8a,
DSDP Hole 47.2, Shatsky Rise, northwest Pacific Ocean];
pl. 133: figs. 1-6 [Zone P8b, DSDP Hole 47.2, Shatsky
Rise, northwest Pacific Ocean]; pl. 137: figs. 2-9 [Zone
P8b, DSDP Hole 20C, South Atlantic Ocean]; pl. 251:
figs. 6 and 7 [Zone of conical globorotaliids, Novogeorgii
Fm., Kuban River section, northern Caucasus].
Globorotalia lensiformis Subbotina subsp. carpatica Samuel,
1972, p. 127-128, pl. 36, figs. 1a-2c (holotype) [middle
Eocene upper Turborotalia (Acarinina) crassata densa
Zone, Myjava brick kiln, north east of Bratislava,
Czechoslovakia].
DESCRIPTION.
Type of wall: Muricate, nonspinose, normal
perforate.
Test morphology: Test low trochospiral,
subquadrate to subcircular, weakly lobulate, chambers
moderately inflated on umbilical side; flat on spiral side
except for initial chambers; surface densely covered by
blunt/truncated muricae giving the test a granular/sugary
texture; 4-41/2 chambers visible in tight coil on umbilical
side; primary aperture a low umbilical-extraumbilical
slit extending to the periphery; sutures on umbilical side
straight to slightly curved, slightly depressed; in spiral
view 9-10 chambers in 21/2-3 whorls, early chambers/
whorls elevated giving biconvex appearance and often
obscured by muricate growth; intercameral sutures
moderately to strongly muricate and (re)curved yielding
trapezoidal shaped chambers; weakly biconvex in edge
view; peripheral muricocarina often obscured by fusion
of muricae along margin.
Size: Diameter: 0.40-0.55 mm; thickness: 0.25-
0.30 mm (Subbotina, 1953, p. 214).
DISTINGUISHING FEATURES.— This taxon is
characterized by its subquadrate, involute, biconvex test
with narrow umbilicus; test covered by moderately to
densely distributed, blunt muricae obscuring, in some
instances, the peripheral muricocarina; 4-41/2 chambers
in last whorl. Later forms exhibit transitional features to
plano-convex, multicameral M. aragonensis.
DISCUSSION.—Subbotina (1953, p. 214) described this
taxon from the lower part of the Zone of conical
BERGGREN AND PEARSON
368
globorotaliids (to which it was said to be essentially
restricted = Zone E4-5 of this paper). She recognized its
descendant affinities with the Globorotalia
marginodentata and G. crassata (=M. aequa-subbotinae
group) and ancestral relationships with M. aragonensis,
interpretations which have withstood the test of time,
relatively unchanged. Shutskaya (1956) subsequently
described the junior synonym Globorotalia nartanensis
from essentially the same stratigraphic level and locality
in the northern Caucasus and recognized its transitional
features with aragonensis.
Blow (1979, p. 981) treated M. dolabrata
(Jenkins) as the ancestor of M. lensiformis (Subbotina)
(Blow, 1979, p. 1005). He distinguished the transition
between the two on the following basis: an increase in
tightness of coiling-mode and proportionate decrease in
size of last chamber relative to earlier chambers and a
relatively stronger recurvature of the spiral intercameral
sutures and more tightly appressed chamber
development in lensiformis. It is clear that Blow (1979)
viewed dolabrata as morphogenetically transitional from
aequa s.s. to lensiformis.
We view the two taxa as synonymous. Topotypes
of dolabrata kindly sent to one of us (WAB) by D.
Graham Jenkins exhibit a densely muricate test with 4-
41/2 chambers as in lensiformis. A (“buried”)
muricocarina rims the test. While Jenkins (1965)
indicated that a peripheral keel was developed only on
the last chamber, his own figures belie this fact (Jenkins,
1965, text-fig. 106), and Blow (1979, p. 401, 982)
pointed out that the presence or visibility of a peripheral
muricocarina is a function of the acuteness of the
peripheral margins of the chambers. In broadly rounded
margins the peripheral muricae are only partially fused
and coalesced and do not yield the same “morphology”
as that seen when the peripheral muricocarinae fuse into
a single band along the margin of a test with an acutely
angled periphery. A buried keel is characteristic of
lensiformis as well and is dependent upon preservation
as well as degree of acuteness of the peripheral margin.
Distinction between these two morphotypes by Blow
(1979) appear to be based on differences of degree rather
than kind and we see little purpose in their separation.
The upper stratigraphic limit of Zone P10 accorded by
Blow (1979, p. 1005) to lensiformis remains enigmatic.
We have not found morphotypes resembling lensiformis
at such stratigraphically high levels.
PHYLOGENETIC RELATIONSHIPS.— This taxon
(probably) evolved from M. subbotinae and is the
ancestor of both M. crater in Zone E4 and M.
aragonensis at the base of Zone E5.
STRATIGRAPHIC RANGE.— Base of Zone E4 to
Zone E6.
GEOGRAPHIC DISTRIBUTION.— Relatively
common in (sub)tropical areas; South Atlantic Ocean,
Indo-Pacific, North Caucasus, among others.
STABLE ISOTOPE PALEOBIOLOGY.— Oxygen and
carbon isotopes indicate a surface mixed layer habitat
(Boersma and others, 1987).
REPOSITORY. Holotype (No. 4111) and paratype
(No. 4112) deposited in the micropaleontological
collections of VNIGRI (Leningrad/St, Petersburg),
Russia.
Morozovella marginodentata (Subbotina, 1953)
PLATE 11.10, FIGURES 1-16
Globorotalia crassata (Cushman).—Glaessner,1937:31, pl.
1: figs. 7a-c [lower Eocene Zone of compressed
globorotaliids, Subzone with Globorotalia
marginodentata, Foraminiferal Beds, Series F1, near
Nal’chik, northern Caucasus, former Soviet Union].—
Subbotina, 1947:119-121 (partim), not pl. 5: figs. 7-12 =
?M. subbotinae (Morozova), pl. 9: figs. 15-17
[Globorotalia ex gr. canariensis, later (1953) designated
the Zone of compressed globorotaliids, Globorotalia
marginodentata Subzone, Kutaiss Horizon (F2),
Kadiizhensk Region, Psish River, northern Caucasus,
former Soviet Union]. [Not Cushman, 1925.]
Globorotalia marginodentata Subbotina, 1953:212, pl. 17:
figs. 14 [lower Eocene, Zone of compressed globorotaliids,
Subzone with Globorotalia marginodentata, Foraminiferal
Beds, Series F1, near Nal’chik, northern Caucasus, former
Soviet Union [refigured from Glaessner, 1937:31, pl. 7:
figs. 7a-c; pl. 17, figs. 15a-c (holotype), and 16a-c, lower
Eocene Zone of compressed globorotaliids, Subzone with
Globorotalia marginodentata, base of Foraminiferal Beds,
Green Series, Kuban River section, northern Caucasus,
former Soviet Union]; pl. 18: figs. 1a-c [lower Eocene
Zone of compressed globorotaliids, Subzone with
Globorotalia marginodentata, Foraminiferal Beds, base
of Kutais Horizon (F2), Khadyzhensk District, Psish River
CHAPTER 11 - MOROZOVELLA
369
section, northern Caucasus, former Soviet Union]; pl. 18:
figs. 2a-c [lower Eocene Zone of conical globorotaliids,
Foraminiferal Beds, Kutai Horizon (F2), Gubs River
section, Barakaevsk District, northern Caucasus, former
Soviet Union]; pl. 18: figs. 3a-c [lower Eocene Zone of
compressed globorotaliids, Foraminiferal Beds, Series F1,
near Nal’chik, Khieu River section, northern Caucasus;
interpreted as transitional form to G. lensiformis].
Luterbacher, 1964:673, text-figs. 75a-76c [lower Eocene
Zone of compressed globorotaliids, Khieu River section,
northern Caucasus, former Soviet Union, determined/
identified by N.N. Subbotina, fide Luterbacher, 1964:674];
77a-78c [uppermost Paleocene Globorotalia velascoensis
Zone, Velasco Fm., Ebano, Mexico; referred to as “aff.
marginodentata”]; 79a-c [lower Eocene G. velascoensis
Zone, Gubbio section, northern Apennines, Italy; referred
to as “aff. marginodentata”]; 80a-c [lower Eocene G.
velascoensis Zone, Gubbio section, Italy]; 81a-82c [lower
Eocene G. aequa Zone, Gubbio section, Italy]; 83a-c
[lower Eocene G. aequa Zone, Gubbio section, Italy;
referred to as “intermediate between Globorotalia
marginodentata and Globorotalia subbotinae”]; 84a-c
[lower Eocene G. formosa formosa/G. subbotinae Zone,
Gubbio section, Italy]. —Samanta, 1970:626, pl. 96: figs.
3,4 [lower Eocene, Pondicherry Fm., Pondicherry, south
east India].—Luterbacher, 1975a:727, pl. 2: figs. 6a-c
[lower Eocene Zone P6, DSDP Hole 313, Mid-Pacific
Mountains].—Luterbacher, 1975b:65, pl. 4: figs. 4-6
[lower Eocene G. subbotinae Zone, Possagno section,
Italy].
Globorotalia (Truncorotalia) aequa marginodentata
Subbotina.—Hillebrandt, 1962:135, pl. 13: figs. 9a-11
[lower Eocene, Zone G, Reichenhall-Salzburg Basin,
Germany].
Truncorotalia marginodentata marginodentata
(Subbotina).—Gohrbandt, 1963:62, pl. 6: figs. 4-6 [lower
Eocene, Zone F, near Salzburg, Austria].
Globorotalia (Morozovella) aequa marginodentata Subbotina.
—Jenkins, 1971:101, text-figs. 177-179 [uppermost
Paleocene G. (S.) triloculinoides Zone, Waipawan Stage,
Middle Waipara River section, North Island, NZ].
Morozovella marginodentata (Subbotina).—Berggren,
1971:76, pl. 5: fig. 9 [lower Eocene Zone P7, DSDP Hole
20C, South Atlantic Ocean].—Berggren, 1977:241, chart
No. 8 [specimens refigured from the literature].—Snyder
and Waters, 1985:460, pl. 8, figs. 13a-14c [lower Eocene
Zone P7, DSDP Hole 549, Goban Spur, Atlantic Ocean].—
Warraich, Ogasawara and Nishi, 2000:293, figs. 17. 4,5,10
[lower Eocene Zone P7, Dungan Fm., Rakhi Nala River,
Sulaiman Range, Pakistan].—Warraich and Ogasawara,
2001:40, fig. 10. 7-9 [lower Eocene Zone P7, Dungan Fm.,
Rakhi Nala River, Sulaiman Range, Pakistan].
Globorotalia (Morozovella) subbotinae (s.l) forma
marginodentata Subbotina.—Blow, 1979:1024-1026, pl.
139: figs. 1-9 and pl. 140: figs.1-3 [lower Eocene Zone
P8b, DSDP Hole 20C, South Atlantic Ocean; see also pl.
223: figs. 5, 6 and pl. 224: figs. 1 and 2].
Truncorotalia marginodentata aperta Gohrbandt, 1963:62,
pl. 5: figs. 10-15 [Zones E and F, near Salzburg, Austria].
DESCRIPTION.
Type of wall: Densely muricate on both sides,
nonspinose, normal perforate.
Test morphology: Low trochospiral, biconvex,
oval to subcircular test with lobulate, thick and strongly
fimbriate muricocarina; chambers on umbilical side
moderately inflated, depressed towards periphery; flat
on spiral side; 41/2-5 chambers in last whorl,
(sub)triangular, compressed along peripheral margin,
rising gradually towards umbilicus; sutures straight to
weakly recurved, radial, depressed; aperture a low,
umbilical-extraumbilical arch or slit extending towards,
but not to, the periphery; umbilicus narrow, deep; in
spiral view approximately 10-12 chambers in 21/2 whorls;
early chambers heavily muricate and elevated above test
surface; chambers lens-shaped to semicircular,
intercameral sutures curved, strongly muricate, elevated
above test surface; scalloped peripheral muricocarinate
periphery characterized by dense concentration of
conical or long slender muricae and the fusion/
coalescence of muricae; in edge view low to moderately
biconvex; early chambers elevated above later whorl(s);
low to medium conical angle (< 45o to ~75o).
Size: Holotype Diameter: 0.43 mm; thickness/
width: 0.24 mm (Subbotina, 1953, p. 213)
DISTINGUISHING FEATURES.— This morphotype,
while exhibiting characters apparently intermediate to
other, closely related forms, is characterized by the
markedly pronounced peripheral compression of the
chambers on the umbilical side which are bordered/
rimmed by a thick and broad peripheral muricocarina
formed by the fusion of inordinately large/thick and long
muricae.
DISCUSSION.— Placement of this taxon is extremely
difficult. “Typical” specimens of marginodentata are
denoted by their marginally compressed/scalloped
chambers and thickly muricate periphery. However, in
most early Eocene (sub)tropical fossil assemblages
intergradation of these morphologies with “typical”
morphologies of M. gracilis and M. subbotinae exist.
Blow (1979, p. 1025, 1026) drew attention to the
BERGGREN AND PEARSON
370
development of marginodentate ornament on both a
subbotinae and gracilis morphotype, interpreted the
marginodentate forms as “extreme phenotypes
developed under particular environmental conditions...”
and observed that “the ‘marginodentate’ forms appear
to occur only in areas of high biotic productivity”. He
chose to view these ecophenotypes nomenclatorially as
extreme forms of subbotinae (sensu stricto) and denoted
them as G. (M.) subbotinae forma marginodentata.
Berggren (1971, p. 76) expressed similar views in
suggesting that marginodentata may be an ecophenotype
of gracilis. We retain the name marginodentata here for
forms possessing the distinctly thick and broad peripheral
muricocarina.
PHYLOGENETIC RELATIONSHIPS.— This
morphotype is closely related to the subbotinae-gracilis
group. It probably evolved from M. subbotinae and does
not appear to have left any descendants.
STRATIGRAPHIC RANGE.— Zone P5 to Zone E5.
GEOGRAPHIC DISTRIBUTION.— Widely distributed
in (sub)tropical regions (Caribbean, Atlantic Ocean,
Indo-Pacific region, North Caucasus)
STABLE ISOTOPE PALEOBIOLOGY.— No data
available.
REPOSITORY.— Holotype (No. 3087) deposited in the
micropaleontological collections at VNIGRI, St
Petersburg, Russia.
Morozovella occlusa (Loeblich and Tappan, 1957)
DISCUSSION. This taxon was discussed in the Atlas
of Paleocene Planktonic Foraminifera (Olsson and
others, 1999).
STRATIGRAPHIC RANGE.— Top of Zone P3b
(typical crosswicksensis); Zone P4 to Zone E2 (typical
occlusa).
Morozovella pasionensis (Bermudéz, 1961)
DISCUSSION.— This taxon was discussed in the Atlas
of Paleocene Planktonic Foraminifera (Olsson and
others, 1999).
STRATIGRAPHIC RANGE.— Zone P3b to Zone E2.
Morozovella subbotinae (Morozova, 1939)
PLATE 11.1, FIGURES 9-16
(Pl. 11.1, Figs. 9-11: new SEMs of paratype of
Globorotalia rex Martin)
Globorotalia subbotinae Morozova, 1939:80, pl. 2: figs. 16-
17 [Paleocene, Emba River, Kazakhstan.].— Luterbacher,
1964:676, text-figs. 85a-c, 86a-c, 89a-c, 90a-c [lower
Eocene G. aequa Zone, Gubbio section, Bottacione Gorge,
northern Apennines, Italy]; figs. 87a-c. [lower Eocene G.
formosa formosa/G. subbotinae Zone, Gubbio section,
Bottacione Gorge, northern Apennines, Italy]; figs. 87a-c
[lower Eocene G. subbotinae Zone, eastern Caucasus,
former Soviet Union].— Proto Decima and Zorzi,
1965:29, 30, pl. 3: figs. 4a-c and pl. 5: fig. 13 [lower
Eocene G. rex Zone, Molinetto di Pederroba section,
Treviso Province, Italy].—Samanta, 1970:635, pl. 96: figs.
1, 2 [G. pseudomenardii /G. subbotinae Zone, Upper
Marlstone, Pondicherry Fm., near Madras, Pondicherry,
South India].—Shutskaya, 1970a:119-120, pl. 13: figs. 6a-
c and pl. 14: figs. 6a-c [lower Eocene G. subbotinae Zone,
Churuk-Su, Bakchissarayan Stage, Kasha River section,
south west Crimea]; pl. 38: figs. 9a-c [G. subbotinae Zone,
Malyi Balkhan Ridge, middle Danatin Fm., western
Turkmenia].—Ferrer, Le Calvez, Luterbacher and Premoli
Silva, 1973:49, text-figs. 11 (2a-c; 5a-c) [G. subbotinae
Zone, Ager Fm., Areny section, Tremp Basin, Catalonia,
Spain].—Stainforth and others, 1975 (partim ):230, text-
fig. 89 (1=holotype refigured), 5-8 (plesiotypes) [G.
subbotinae Zone, Lodo Fm., Fresno Co., California]; figs.
3a-c [from Subbotina, 1953, pl. 17: figs. 13a-c refigured,
as G. crassata Cushman]; not fig. 89 (2a-c) [from
Subbotina, 1953, pl. 17: figs. 11a-c (refigured) as G.
crassata Cushman=M. aequa (Cushman and Renz)].—
Luterbacher, 1975a:72, pl. 2: figs. 2a, b [G. formosa
formosa Zone, DSDP Hole 305, Shatsky Rise, north-west
Pacific Ocean].—Luterbacher, 1975b:65, pl. 2: figs. 31-
33 [Possagno section, Italy].—Murray and others,
1989:532, pl. 10.10: fig. 31 [lower Eocene Zone NP 11,
Bracklesham Group, English Channel]; pl. 10.11, fig. 1
[lower Eocene Wittering Fm., East Wittering, Sussex, UK].
Globorotalia (Morozovella ) subbotinae subbotinae
(Morozova).—Blow, 1979:1018-1021, pl. 102: figs. 1-5
and pl. 219: figs. 1-4 [Zone P6 , DSDP Hole 20C, South
Atlantic Ocean]; pl. 111: figs. 6-8 [Zone P7, Moogli
Mudstones, Kagua, Papua, New Guinea]; pl. 115: figs. 3-
5 [Zone P7, Kilwa area, Tanzania]; also pl. 115: fig. 6 as
Globorotalia (Morozovella) sp. specimen ex interc G. (M.)
aequa lacerti Cushman and Renz and G. (M.) subbotinae
subbotinae Morozova [lower Eocene, Zone P7, Kilwa
area, Tanzania]; pl. 119: figs. 4-10; pl. 219: figs. 5, 6; pl.
220: figs. 1-6 and pl. 222: figs. 1-7 [Zone P7 , DSDP Hole
CHAPTER 11 - MOROZOVELLA
371
47.2, Shatsky Rise, northwest Pacific Ocean]; pl. 127:
figs. 6, 7 [Zone P8a, DSDP Hole 47.2, Shatsky Rise, north-
west Pacific Ocean]; pl. 123: fig. 8 [Zone P8b, DSDP Hole
47.2/, Shatsky Rise, northwest Pacific Ocean]; pl. 103:
figs. 2, 7, 9 and pl. 223: figs. 1, 2 [Zone P6, DSDP Hole
20C, South Atlantic Ocean).
Globorotalia crassata (Cushman).—Subbotina, 1947
(partim):119, pl. 5: figs. 31-33 [lower Eocene Globorotalia
ex. gr. canariensis Zone of Subbotina, 1947 = G. crassata
Subzone of Zone of compressed globorotaliids of
Subbotina, 1953, Assa River, northern Caucasus]; not pl.
9: figs. 15-17 [lower Eocene G. ex. gr. canariensis Zone,
Khieu River section, northern Caucasus; included in
synonymy of Globorotalia marginodentata Subbotina,
1953].—Subbotina, 1953 (partim):211, 212, pl. 17: figs.
7a-c (holotype of G. subbotinae Morozova refigured); pl.
17, figs. 13a-c [Zone of compressed globorotaliids,
Tarkhankut Peninsula, Crimea].
Morozovella subbotinae (Morozova).—Berggren, 1971:76 pl.
5: figs. 10, 11 [G. subbotinae Zone, DSDP Hole 20C, South
Atlantic Ocean].—Berggren, 1977:239, 240 (illustrations
refigured from literature).—Snyder and Waters, 1985:442,
443, pl. 9: figs. 10-12 [Zone P6, DSDP Hole 548A, Goban
Spur, northeast Atlantic Ocean].—Toumarkine and
Luterbacher, 1985:112, text-figs. 15: 9a-c (reillustration
of holotype of G. rex Martin, 1943); 10a-c (reillustration
of holotype of G. subbotinae Morozova, 1939); 11a-c
(reillustration of specimen from northwest Crimea
identified by Subbotina, 1953, pl. 17: figs. 13a-c as G.
crassata Cushman ).—Huber, 1991:440, pl. 4, fig. 9 [Zone
AP6A, ODP Hole 738, southern edge of Kerguelen
Plateau, South Indian Ocean].—Lu and Keller, 1993:123,
pl. 4: fig. 19 [lower Eocene M. subbotinae Subzone, ODP
Hole 738C, southern edge of Kerguelen Plateau, South
Indian Ocean].—Lu and Keller, 1995:102, pl. 1: figs. 11-
13 [Subzone P6b, DSDP Hole 577, Shatsky Rise, north-
west Pacific Ocean].—Basov, 1995:165, pl. 2: figs. 8-10
[M. subbotinae Zone, ODP Hole 883B, Detroit Seamount,
northwest Pacific Ocean).—Bralower and others,
1995:844, fig. 2 (3,4) [Zone P4/5, ODP Hole 865C, Allison
Guyot, equatorial west Pacific Ocean].—Warraich and
Ogasawara, 2001:41, figs. 10.16-18 [Zone P7, Dungan
Fm., eastern limb of Zinda Pir anticline, Sulaiman Range,
Pakistan].
Globorotalia rex Martin, 1943:117, pl. 8: figs. 2a-c [lower
Eocene Zone P6, Lodo Fm., Lodo Gulch, Fresno Co.,
California].—Bolli, 1957b:75, pl. 18: figs. 10-12 [G. rex
Zone, Upper Lizard Springs Fm., Trinidad].—Postuma,
1971:210, figs. on p. 211 [topotypes from G. rex Zone,
locality no. M-74, Lodo Fm., Fresno Co., California].—
Said and Sabry, 1964:385, pl. 2: figs. 4a-c [G. rex Zone,
Upper Esna Shale, Gebel Owaina, Egypt].
Truncorotalia cf. rex (Martin) .—Gohrbandt, 1963:64, pl. 6:
figs. 1-3 [Zone F, Salzburg Basin, Austria].
Globorotalia (Morozovella) aequa rex Martin.—Jenkins,
1971:101, 102, pl. 7: figs. 180-182 [G. wilcoxensis Zone,
Waipawan Stage, Middle Waipara River section, NZ].
Globorotalia bollii El Naggar, 1966 (partim, not pl. 22: figs.
6a-c [same level; = M. gracilis (Bolli]):202, 203, pl. 22:
figs. 5a-d [holotype from G. wilcoxensis Zone, Thebes
Limestone, Gebel Owaina section, Egypt].
Globorotalia nartanensis Shutskaya.—Shutskaya 1970b:118-
120, pl. 15: figs. 2a-c and 8a-c [G. subbotinae Zone,
Churuk-Su, Bakhchissarayan Stage, Kacha River section,
Bakhchissaray region, south west Crimea]. [Not
Shutskaya, 1956.]
Globorotalia (Truncorotalia) aequa simulatilis Schwager.—
Hillebrandt, 1962:134, pl. 13: figs. 6a-c; 7, 8a-c [Zone G,
Reichenhall-Salzburg Basin, Germany and Austria]. [Not
Schwager, 1883.]
Not Globorotalia rex Martin.—Loeblich and Tappan,
1957:195, pl. 60: figs., 1a-c [G. pseudomenardii Zone,
Nanafalia Fm., Wilcox Co., Alabama; ?=M. velascoensis
parva ; ?= M. acuta ].—Mallory, 1959:40, pl. 34 figs. 12a-
c; pl. 42 figs. 2a-c [Ynezian-Narizian of California Coast
Ranges = M. aragonensis (Nuttall)].
DESCRIPTION.
Type of wall: Normal perforate, muricate,
nonspinose.
Test morphology: Test relatively large (to 0. 5 mm
maximum diameter), planoconvex to weakly biconvex
test with moderately lobulate, strongly/thickly keeled
periphery; 4-41/2 chambers in last whorl, generally
covered with muricae on umbilical side, spiral side
relatively smooth; umbilical and spiral intercameral
sutures weakly curved, tangential on spiral side yielding
trapezoidal- shaped chambers; circumumbilical chamber
tips weakly ornamented by muricae and surrounding
deep, narrowly open umbilicus; aperture a low,
umbilical-extraumbilical slit extending almost to
periphery and bordered by weak lip.
Size: Holotype diameter: 0.36 mm; thickness: 0.20
mm.
DISTINGUISHING FEATURES.— Morozovella
subbotinae is distinguished by its relatively large and
strongly muricocarinate test, and the circumumbilical
elevation/extension of the chambers and weakly
ornamented circumumbilical chamber tips.
BERGGREN AND PEARSON
372
DISCUSSION. This robust morphospecies is a
characteristic element of latest Paleocene and early
Eocene planktonic foraminiferal assemblages.
Morozovella subbotinae has had a convoluted taxonomic
history. It is generally agreed by specialists that
Globorotalia subbotinae Morozova, 1939 is a senior
synonym of Globorotalia rex Martin, 1943 (see
Berggren, 1977; Blow, 1979 for discussions). In the
(former) Soviet Union this taxon was identified with (the
middle Eocene) Globorotalia crassata Cushman, 1925
(see our treatment in Pearson and Berggren, Chapter 10,
this volume; see also Subbotina, 1947, p. 119-121; 1, p.
211). Included in this taxon subsequently by Subbotina
(1953) were forms referable to M. aequa and M.
subbotinae (including the holotype reference of
Morozova, 1939, of G. subbotinae) as well as forms
(1947, pl. 9, figs. 15-17) subsequently (1953, pl. 18, figs.
1a-c) referred to the new taxon Globorotalia
marginodentata Subbotina, 1953. Examination of a suite
of specimens identified as G. crassata donated to one
of us (WAB) in 1962 by Subbotina bears this out.
An enigmatic case of taxonomic affinities is
provided by Globorotalia nartanensis Shutskaya, 1956.
The (relatively poor) illustration of the holotype is
balanced by a clear and thorough description of the
taxon. The low, biconvex test, thick, blunt “spines”
(=muricae) on the test surface which give the test the
appearance of having a thick, “granular” test wall and
the transitional characters to M. aragonensis noted by
Shutskaya (1956) make it quite clear that this taxon, in
its original concept, is a junior synonym of Globorotalia
lensiformis Subbotina, 1953. However, Shutskaya
(1972b) subsequently illustrated two morphotypes
identified as G. nartanensis from the G. subbotinae Zone
of the south west Crimea which are virtually identical
with Subbotina’s (1953, pl. 17, figs. 13a-c) illustrations
of a peripherally compressed test with development of
a frilled (“marginodentate”) keel which she considered
transitional between Globorotalia crassata (=G. aequa/
subbotinae) and G. marginodentata.
Blow (1979, p. 1018-1026) has drawn attention to
the close similarities between M. subbotinae
(Morozova), M. marginodentata (Subbotina) and M.
gracilis (Bolli). In fact he considered gracilis separated
from subbotinae at the subspecies level based on the
increase in chamber number (from 41/2 in subbotinae to
51/2-6 in gracilis), associated with the development of a
somewhat more evolute coiling-mode and more
vorticiform spiral intercameral sutures in gracilis and,
finally, the slightly different (shorter) stratigraphic range
of gracilis. The development of a strongly dentate
(fimbriate) muricocarina (marginodentata) on some
morphotypes was considered little more than
ecophenotypic variation within the subbotinae plexus
of morphotypes, indicative of high productivity, and
marginodentata was, accordingly, considered only a
variant of subbotinae (cf. Berggren, 1971, who had
suggested earlier that marginodentata might be
synonymous with, or an ecophenotypic variant of,
gracilis).
PHYLOGENETIC RELATIONSHIPS. This taxon
evolved from M. aequa by an increase in test size,
developing a thicker, broader peripheral muricocarina
and more pronounced angulo-conical test; it evolved into
M. lensiformis and also gave rise to M. marginodentata
and M. gracilis.
STRATIGRAPHIC RANGE.Zone P5 to Zone E5. We
have found that M. subbotinae has its FAD at the top of
Chron C25n at ODP Site 577 and has a short overlap in
the upper part of its range with Morozovella aragonensis.
Its LAD is used to denote the boundary between Zones
E5 and E6 here. The premature disappearance of M.
subbotinae (within Zone P6b = E4) at Indian Ocean Site
213 is ascribed to gradually increasing dissolution in
the early Eocene.
GEOGRAPHIC DISTRIBUTION. Widely distributed
in (sub)tropical assemblages in Atlantic, Indo-Pacific,
and typical Tethyan biogeographies and as far south as
60o S in association with the early Eocene extra-tropical
excursion of carinate morozovellids on the Kerguelen
Plateau (Huber, 1991, ODP Site 738; Berggren, 1992,
ODP Site 747; see also Olsson and others, 1999, p. 67,
text-figure 24).
STABLE ISOTOPE PALEOBIOLOGY. Morozovella
subbotinae has δ13C and δ18O values similar to M.
velascoensis and Acarinina nitida and has more positive
δ13C and δ18O than Subbotina triangularis (D’Hondt and
others, 1994). Morozovella subbotinae displays a
pronounced increase in δ13C with increased test size but
little corresponding change in δ18O (D’Hondt and others,
1994).
CHAPTER 11 - MOROZOVELLA
373
REPOSITORY. Holotype (slide no. 700) deposited
in the micropaleontological collections of VNIGRI, St.
Petersburg, Russia.
Morozovella velascoensis (Cushman, 1925)
PLATE 11.2, FIGURES 1-3
DISCUSSION.— This taxon was discussed in the Atlas
of Paleocene Planktonic Foraminifera (Olsson and
others, 1999).
STRATIGRAPHIC RANGE.— P3b to Zone E2.
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376
... They also do not illustrate the transitional specimens which document the origin and phylogenetic relationships of Ac. sibaiyaensis proposed above. The authors also neglected the considerable difference in size of Ac. africana (Berggren et al., 2006a) and M. allisonensis (Berggren and Pearson, 2006) by unifying the size of the images. Guasti and Speijer (2008), on the other hand, stated that Ac. africana and Ac. ...
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