GeolCarp_Vol64_No3_237

www.geologicacarpathica.sk

GEOLOGICA CARPATHICA

, JUNE 2013, 64, 3, 237—251 doi: 10.2478/geoca-2013-0016

Introduction

After  the  Cretaceous/Paleogene  (K/Pg)  boundary  mass  ex-
tinction event a planktonic foraminiferal evolutionary radia-
tion  led  to  the  origin  to  some  of  the  main  Early  Paleogene
lineages (Premoli Silva 1977; Smit 1982; D’Hondt 1991; Liu
&  Olsson  1992;  MacLeod  1993;  Olsson  et  al.  1999;
Apellániz  et  al.  2002;  Arenillas  et  al.  2010).  One  of  these
leads to the genus Globanomalina Haque, 1956, and another
to  Praemurica  Olsson,  Hemleben,  Berggren  &  Liu,  1992.
They  were  designated  “non-spinose  lineages”  by  Olsson  et
al.  (1999).  Both  exhibit  > 1 µm-diameter  pore-pits  (also
called  pore-funnels).  These  are  pores  ending  in  a  funnel-
shaped  depression  in  the  external  test  wall  enlarging  exter-
nally  the  pore.  However,  while  Praemurica  develops
honeycomb-like surface ridges (interpore ridges) typical of a
cancellate  or  reticulate  wall,  Globanomalina  exhibits  a  less
ornamented  wall,  so  its  texture  has  been  traditionally  de-
scribed as smooth. None of them have spines, papillas and/
or pustules.

The phylogenetic origin of these genera has been the sub-

ject  of  several  studies,  in  particular  those  of  Olsson  et  al.
(1992, 1999) and Liu & Olsson (1994). They supported the
previous  hypotheses  of  Berggren  (1962,  1977),  Olsson
(1963,  1970)  and  Blow  (1979),  suggesting  that  the  normal
perforate Danian planktonic foraminifera (i.e. pitted and can-
cellate  walls)  derived  from  Hedbergella  Brönnimann  &
Brown, 1958 with pitted walls. According to them, the can-
cellate  (spinose  and  non-spinose)  species  derived  from  H.
monmouthensis  Olsson,  1960,  and  the  pitted-smooth  one
from H. holmdelensis Olsson, 1964. In contrast, Arenillas &

New evidence on the origin of non-spinose pitted—cancellate

species of the early Danian planktonic foraminifera

IGNACIO ARENILLAS and JOSE ANTONIO ARZ

Departamento de Ciencias de la Tierra, and Instituto Universitario de Investigación en Ciencias Ambientales de Aragón, Universidad de

Zaragoza, E-50009 Zaragoza, Spain; ias@unizar.es; josearz@unizar.es

(Manuscript received July 24, 2012; accepted in revised form December 11, 2012)

Abstract: Intermediate forms identified in some of the most continuous lower Danian sections allow a better understand-
ing of the origin and evolution of pitted (Globanomalina) and cancellate (Praemurica) planktonic foraminifera. Both
Globanomalina and Praemurica are part of a major Paleocene lineage, namely the “non-spinose lineage”, which started to
diverge in the early Danian. Transitional specimens strongly suggest the evolution from Parvularugoglobigerina to
Globanomalina, and then to Praemurica. These evolutionary turnovers were quite rapid (probably lasting less than 10 kyr),
and seem to have begun in the time equivalent of the lower part of the E. simplicissima Subzone, namely the middle part of
the standard Zone Pa. The initial evolutionary trends within this non-spinose lineage were the increase of test size and lip
thickness, and the evolution from tiny pore-murals to large pore-pits, and from smooth to pitted and finally cancellate
walls. Biostratigraphic data suggest that evolution of the wall texture preceded the morphological evolution within each
genus. The oldest species of both Globanomalina and Praemurica, namely G. archeocompressa and Pr. taurica, initially
retained the external morphology of the ancestral Parvularugoglobigerina eugubina. Since their divergence, Globanomalina
and Praemurica followed a separate evolutionary path, evolving into morphologically different species.

Key words: Paleocene, phylogeny, planktonic foraminifera, wall texture, Parvularugoglobigerina, Globanomalina,
Praemurica.

Arz (1996, 2000) and Arenillas et al. (2010) suggested the
alternative perspective that Palaeoglobigerina Arenillas, Arz
& Nán

~ez, 2007, was the ancestor of the spinose lineage,

whereas Parvularugoglobigerina Hofker, 1978 was the fore-
runner of the non-spinose lineage, first giving rise to Globa-
nomalina  and  then  to  Praemurica.  A  similar  phylogenetic
hypothesis  had  already  been  proposed  by  Premoli  Silva
(1977),  who  considered  that  Globanomalina,  in  particular
Globanomalina  compressa  (Plummer,  1927)  derived  from
Parvularugoglobigerina,  specifically  from  Parvularugoglo-
bigerina eugubina (Luterbacher & Premoli Silva, 1964).

Following an intensive search of transitional lowermost

Danian specimens, this study proposes an alternative phylo-
genetic  hypothesis  on  the  origin  of  the  non-spinose  lineage,
suggesting an evolutionary relationship between smooth Par-
vularugoglobigerina and pitted Globanomalina, and between
the latter and the cancellate Praemurica. In addition, we pro-
pose phylogenetic relationships among species based on tex-
tural  and  external  morphological  criteria  and  high  resolution
biostratigraphic data. In order to carry out this study, we have
used  specimens  from  some  of  the  most  continuous  and  ex-
panded stratigraphic sections available from the lower Danian.

Material and methods

For the analysis of the biostratigraphic ranges of the stud-

ied taxa and their transitional specimens, we have revised the
lower Danian in the El Kef and A

n Settara sections (Tunisia)

and in the Caravaca and Agost sections (Spain). We also
have taken into account biostratigraphic data from other sec-

238

ARENILLAS and ARZ

GEOLOGICA CARPATHICA

GEOLOGICA CARPATHICA, 2013, 64, 3, 237—251

tions  such  as  Elles  (Tunisia),  Ben  Gurion  (Israel),  Gubbio
(Italy),  Zumaia,  Osinaga  and  San  Sebastián  (Spain),  Bidart
(France),  El  Mulato,  El  Mimbral,  La  Lajilla,  Bochil  and
Guayal (Mexico), and Loma Capiro (Cuba). The geographi-
cal location of the analysed sections is shown in Fig. 1.

We used the lower Danian planktonic foraminiferal zona-

tions  by  Arenillas  et  al.  (2004)  and  Berggren  &  Pearson
(2005); their equivalence is shown in Fig. 2. The zonation of
Arenillas et al. (2004) includes the three classical lower Danian
Guembelitria  cretacea,  Parvularugoglobigerina  eugubina
and  Parasubbotina  pseudobulloides  Zones,  and  divides  each
of these biozones into two Subzones: Hedbergella holmdelen-
sis  and  Parvularugoglobigerina  longiapertura  Subzones  for
the  G.  cretacea  Zone,  Parvularugoglobigerina  sabina  and
Eoglobigerina  simplicissima  Subzones  for  the  Pv.  eugubina
Zone,  and  Eoglobigerina  trivialis  and  Subbotina  triloculi-
noides  Subzones  for  P.  pseudobulloides  Zone.  We  have  in-
cluded in this paper a third subzone in the upper part of the
P. pseudobulloides Zone: the Globanomalina compressa Sub-
zone, which was given the rank of Zone by Arenillas & Molina
(1997).  As  shown  in  Fig. 2,  Berggren  &  Pearson’s  (2005)
Zone  P0  is  equivalent  to  the  H.  holmdelensis  Subzone,  the
Zone Pa approximately spans both the P. longiapertura Sub-
zone and the Pv. eugubina Zone, and P1a, P1b and P1c are

Fig. 1. Geographical location of the lower Danian sections mentioned
in the text: 1 – El Kef, Aïn Settara and Elles (Tunisia); 2 – Cara-
vaca and Agost (S Spain); 3 – Zumaia, Osinaga and San Sebastián
(N Spain); 4 – Bidart (France); 5 – Gubbio (Italy); 6 – Ben Gurion
(Israel);  7 – El Mulato, El Mimbral and La Lajilla (NE Mexico);
8 – Bochil and Guayal (S Mexico);  9  –  Loma  Campiro  (Cuba);
10  –  Lynn  Creek  (Mississippi);  11  –  Bajada  del  Jagüel  (Argen-
tine); 12 – DSDP Site 305 (North Pacific).

Fig. 2. Biostratigraphic ranges of analysed species: 1 – planktonic foraminiferal zo-
nation of Arenillas et al. (2004 modified); 2 – planktonic foraminiferal zonation of
Berggren & Pearson (2005); dotted lines mean doubtful biostratigraphic distribution
(not supported by SEM-photographed specimens).

roughly equivalent to the E. trivialis, S.
triloculinoides and G. compressa Subzones
respectively. The ranges of species shown
in Fig. 2 are based on biostratigraphic data
from the sections mentioned above.

For morphological analysis, we also

checked  other  sections  and  boreholes
(Fig. 1) such as Elles (Tunisia), Ben Gurion
(Israel),  Bajada  del  Jagüel  (Argentine),
Lynn  Creek  (Mississippi)  and  DSDP
Site 305  (Shatsky  Rise,  North  Pacific).
Specimens  were  chosen  mainly  from  El
Kef  and  Aïn  Settara  samples,  which  were
disaggregated  in  water  with  diluted  H

and  washed  through  a  63-µm  sieve.  The
foraminiferal preservation in these sections
is good enough to analyse the wall texture,
although  corroded  and  recrystallized  sur-
faces  can  be  observed  in  some  specimens
(“frosty”  specimens  according  to  the
terminology  of  Sexton  et  al.  2006).  Wall
textures  were  examined  under  scanning
electron  microscopes  (SEM),  trademarks
JEOL  JSM  6400  and  Zeiss  MERLIN
FE-SEM, at the Electron Microscopy Ser-
vice  of  the  Universidad  de  Zaragoza
(Spain).  Over  600  SEM-photographs,  in-
cluding  different  views  of  whole  speci-
mens, were taken of 170 specimens, some
of which are morphotypes transitional be-
tween  species.  Except  for  type-specimens
of  other  authors,  all  the  specimens  illus-
trated  in  Figs. 3  to  8  are  deposited  in  the
Departamento  de  Ciencias  de  la  Tierra  of
the Universidad de Zaragoza (Spain).

239

EARLY DANIAN NON-SPINOSE PITTED—CANCELLATE PLANKTONIC FORAMINIFERA

GEOLOGICA CARPATHICA

GEOLOGICA CARPATHICA, 2013, 64, 3, 237—251

Taxonomic and phylogenetic notes

The taxonomy used here is based mainly on that of Arenil-

las  (1996),  with  some  modifications.  The  species  analysed
were: Hedbergella monmouthensis Olsson (1960; Fig. 3A—B),
Hedbergella holmdelensis Olsson, 1964 (Fig. 3C—I), Parvu-
larugoglobigerina  longiapertura    (Blow  1979;  Fig. 3J—K),
Parvularugoglobigerina  eugubina  (Luterbacher  &  Premoli
Silva  1964;  Fig. 3L—O),  Globanomalina  archeocompressa
(Blow 1979; Fig. 4A—F), Globanomalina imitata (Subbotina
1953; Fig. 4G—J), Globanomalina planocompressa (Shutskaya
1965;  Fig. 4K—4N),  Globanomalina  compressa  (Plummer
1927;  Fig. 5A—B),  Praemurica  taurica  (Morozova  1961;
Fig. 5C—D),  Praemurica  pseudoinconstans  (Blow  1979;
Fig. 5E—G),  and  Praemurica  inconstans  (Subbotina  1953;
Fig. 5H—L).  The  diagnostic  characteristics  of  these  species
are given in Appendix.

Wall texture has been widely considered when analysing

the taxonomy and phylogeny of planktonic foraminifera. It is
allowing  taxa  to  be  distinguished  at  the  genus  rank  (e.g.
Nederbragt  1991;  Olsson  et  al.  1992;  Liu  &  Olsson  1992,
1994; Georgescu 2007, 2009a,b; Arenillas et al. 2010). Olsson
et al. (1999) differentiated two major lineages in the Danian
based on textural criteria: the “non-spinose lineage” (Globa-
nomalina and Praemurica) and the “spinose lineage” (Eoglo-
bigerina  Morozova,  1959,  Subbotina  Brotzen  &  Pozaryska,
1961,  and  Parasubbotina  Olsson,  Hemleben,  Berggren  &
Liu, 1992). They also proposed two lineages within the lower
Danian non-spinose: “smooth-walled lineage” (Globanoma-
lina)  and  “praemuricate  lineage”  (Praemurica).  These  lin-
eages  are  referred  to  here  as  “non-spinose  pitted”  and
“non-spinose  cancellate”  respectively  because  Globanomalina
exhibits a pitted rather than smooth wall texture (i.e. its pore-
pits are always larger than 1 µm in diameter, and can easily
be  observed  under  the  stereomicroscope),  and  the  term
“praemuricate”  is  not  descriptive  enough.  Defined  in  this
way,  the  Globanomalina  wall  texture  can  be  distinguished
from  the  smooth  wall  of  Parvularugoglobigerina  and  the
non-spinose cancellate wall of Praemurica.

Blow (1979) ascribed non-spinose pitted and cancellate

Early Paleogene taxa to the large traditional family Globoro-
taliidae  Cushman,  1927,  specifically  to  Globorotalia  (Tur-
borotalia)  Cushman  &  Bermúdez,  1949.  He  suggested  that
Hedbergella was the ancestor of these early “turborotaliids”,
based  mainly  on  similarities  with  the  Danian  pseudobul-
loides-group  (currently  separated  from  the  non-spinose  lin-
eage and assigned to Parasubbotina by Olsson et al. 1999).
Blow  (1979)  proposed  that  some  earliest  Danian  species
with  smooth  wall  texture,  such  as  Pv.  longiapertura  (Blow
1979;  Fig. 3J—K),  should  be  included  in  Globorotalia  (Tur-
borotalia).  However,  this  group  of  primitive  species  had
been assigned previously to the earliest Danian genus Parvu-
larugoglobigerina  Hofker,  1978.  Olsson  et  al.  (1999)  ex-
cluded  Parvularugoglobigerina  from  Globorotaliidae,  after
suggesting close relationships between parvularugoglobiger-
inids and guembelitriids and ascribed it to the family Guem-
belitriidae Montanaro Gallitelli, 1957.

Loeblich & Tappan (1987) classified Globanomalina in the

family Globanomalinidae Loeblich & Tappan, 1984, which

was  defined  as  comprising  Paleogene  species  with  non-spi-
nose  “smooth”  wall  and  low  trochospiral  to  planispiral  coil-
ing.  This  macrotaxonomic  classification  was  adopted  by
Arenillas  (1996).  However,  Olsson  et  al.  (1999)  concluded
that Globanomalina was derived from the Late Maastrichtian
hedbergellids,  and  consequently  included  it  in  the  family
Hedbergellidae Loeblich & Tappan, 1961. The wall of Hed-
bergella is similar to that of Globanomalina, namely pitted—
smooth,  but  displays  small  scattered  pustules,  which  are
frequently  very  abundant  over  the  first  chambers  of  the  test
(Fig. 3A—I).  Olsson  et  al.  (1999)  and  Apellániz  et  al.  (2002)
suggested H. holmdelensis was the ancestral species of Globa-
nomalina,  the  former  Danian  species  being  G.  archeocom-
pressa.  According  to  this  hypothesis,  the  morphological
characters of Globanomalina species (compressed test with an
imperforate peripheral band, and an umbilical—extraumbilical
aperture bordered by a narrow lip) were derived directly from
the ancestral species.

Praemurica, whose type-species is Pr. taurica, was de-

fined  in  order  to  group  Lower  Paleocene  species  with  low
trochospiral and well-developed cancellate wall texture (Olsson
et  al.  1992).  It  was  the  first  non-spinose,  cancellate  taxa  to
evolve  after  the  K/Pg  boundary  mass  extinction  (Olsson  et
al. 1999). Others authors (e.g. Arenillas 1996) included it in
the  family  Truncorotaloididae  Loeblich  &  Tappan,  1961.
Olsson et al. (1999) also suggested a hedbergellid origin for
Praemurica,  but  independently  of  the  Globanomalina  ori-
gin,  proposing  H.  monmouthensis  as  the  ancestral  species,
and Pr. taurica as its first species to appear. A similar phylo-
genetic scenario was proposed by Apellániz et al. (2002), but
they suggested that Praemurica derived from H. holmdelen-
sis, with Pv. eugubina as an intermediary step.

Arenillas & Arz’s (1996) hypothesis proposed alternatively

that  Parvularugoglobigerina  was  the  ancestor  of  the  entire
non-spinose  lineage  and  the  genus  Globanomalina  the  first
one  to  evolve.  Arenillas  (1996)  therefore  ascribed  Parvu-
larugoglobigerina  to  the  family  Globanomalinidae  together
with  Globanomalina.  These  authors  also  suggested  that
Praemurica  evolved  somewhat  later  deriving  from  G.  ar-
cheocompressa, with Pr. taurica being its first species. Ac-
cording  to  this  hypothesis,  Parvularugoglobigerina  had
previously derived from Hedbergella, in particular from ear-
liest Danian

dwarfed H. holmdelensis, based on morphologi-

cal similarities and biostratigraphic data. However, a more
detailed morphological and textural analysis (Arenillas et al.
2007, 2010) indicated that Parvularugoglobigerina derived
from Palaeoglobigerina, which in turn evolved from Guembe-
litria Cushman, 1933.

Olsson et al. (1999) and Arenillas et al. (2007) disagree on

the wall structure of the parvularugoglobigerinids. This first
group  of  authors  claims  that  parvularugoglobigerinids  have
an  irregularly  pore-mounded  wall;  the  second  group  main-
tains that their wall is smooth. The discrepancy is not caused
by  poor  preservation,  since  smooth  walled  specimens  and
specimens  with  more  ornamented  walls  (i.e.  pore-mounded
Guembelitria, and/or rugose Woodringina) are found in the
same  samples  and  exposed  to  the  same  diagenetic  condi-
tions (Arenillas et al. 2010). Recently, Arenillas et al. (2012)
have suggested the co-occurrence of two groups of primitive

240

ARENILLAS and ARZ

GEOLOGICA CARPATHICA

GEOLOGICA CARPATHICA, 2013, 64, 3, 237—251

Fig. 3. SEM images of Hedbergella Brönnimann & Brown, 1958, and Parvularugoglobigerina Hofker, 1978, species of the K-Pg transition
(scale bar = 100 µm; scale bar of SEM-micrographs illustrating details = 10 µm). A – Hedbergella monmouthensis Olsson, 1960, topotypes
from Olsson et al. (1999), from the upper Maastrichtian, Redbank Fm, New Jersey: 1 – umbilical view, 2 – axial view. B – H. mon-
mouthensis, from the H. holmdelensis Subzone (G. cretacea Zone), Aïn Settara, Tunisia: 1 – spiral view, 2 – axial view, 3 – umbilical view.
C, D – Hedbergella holmdelensis Olsson, 1964, two topotypes of Olsson et al. (1999), from the upper Maastrichtian, Redbank Fm, New
Jersey: C – umbilical view, D – axial view. E – H. holmdelensis, from the Pv. longiapertura Subzone (G. cretacea Zone), Aïn Settara,
Tunisia: 1 – umbilical view, 2 – axial view, 3 – spiral view, 4, 5 – details of surface. F – H. holmdelensis, from the H. holmdelensis Sub-
zone (G. cretacea Zone), Aïn Settara, Tunisia: 1 – umbilical view, 2 – axial view, 3 – spiral view. G, H, I – H. holmdelensis, three
specimens from Olsson et al. (1999) assigned to Globanomalina archeocompressa (Blow, 1979), from the Zone P0, Millers Ferry, Alabama:
G – umbilical view, H – axial view, I – spiral view. J – Parvularugoglobigerina longiapertura (Blow, 1978), from the Pv. longiapertura
Subzone (G. cretacea Zone), Aïn Settara, Tunisia: 1 – umbilical view, 2 – axial view. K – Pv. longiapertura, type-specimens from Blow
(1979), from the Zone Pa, DSDP Leg 6, South Pacific: 1 – holotype, umbilical view, 2 – paratype, axial view. L – Parvularugoglobigerina
eugubina (Luterbacher & Premoli Silva, 1964), “Pv. eugubina” type-sample, from the lowermost Danian, Ceselli, Italy: 1 – spiral view,
2 – axial view, 3 – umbilical view. M – Pv. eugubina, from the E. simplicissima Subzone (Pv. eugubina Zone), Aïn Settara, Tunisia:
1 – axial view, 2 – umbilical view. N – Pv. eugubina (Luterbacher & Premoli Silva), from the E. simplicissima Subzone (Pv. eugubina
Zone), Agost, Spain: 1 – umbilical view, 2 – axial view. O – Pv. eugubina, from the Pv. sabina Subzone (Pv. eugubina Zone), El Kef,
Tunisia: 1 – spiral view, 2 – axial view, 3 – umbilical view, 4 – detail of surface (granular wall texture in parvularugoglobigerinids).

241

EARLY DANIAN NON-SPINOSE PITTED—CANCELLATE PLANKTONIC FORAMINIFERA

GEOLOGICA CARPATHICA

GEOLOGICA CARPATHICA, 2013, 64, 3, 237—251

Fig. 4. SEM images of Globanomalina Haque (1956) species of the lower Danian (scale bar=100 µm). A – Globanomalina archeocom-
pressa (Blow, 1979), paratype, Zone Pa, DSDP Leg 6, South Pacific (Blow 1979): umbilical view. B – G. archeocompressa, holotype,
from the Zone Pa, DSDP Leg 6, South Pacific: umbilical view. C – G. archeocompressa, paratype, from Zone Pa, DSDP Leg6, South Pa-
cific: axial view. D – G. archeocompressa, from the S. triloculinoides Subzone (P. pseudobulloides Zone), El Kef, Tunisia: 1 – umbilical
view, 2 – axial view, 3 – spiral view. E – G. archeocompressa, from the E. trivialis Subzone (P. pseudobulloides Zone), Aïn Settara,
Tunisia: 1 – spiral view, 2 – umbilical view. F – G. archeocompressa, from the G. compressa Subzone (P. pseudobulloides Zone), Caravaca,
Spain: 1 – umbilical view, 2 – axial view. G – Globanomalina imitata (Subbotina, 1953), holotype, from Danian, Kuban River, northern
Caucasus, Russia (SEM-image from Olsson et al. 1999): 1 – spiral view, 2 – axial view, 3 – umbilical view. H – G. imitata, from the S.
triloculinoides Subzone (P. pseudobulloides Zone), El Kef, Tunisia: 1 – umbilical view, 2 – axial view, 3 – spiral view. I – G. imitata
(Subbotina), from the E. trivialis Subzone (P. pseudobulloides Zone), El Kef, Tunisia: 1 – spiral view, 2 – axial view, 3 – umbilical view.
J – G. imitata, transitional to G. planocompressa, from the E. trivialis Subzone (P. pseudobulloides Zone), Osinaga, Spain: 1 – axial view,
2 – umbilical view. K – Globanomalina planocompressa (Shutskaya, 1965), holotype, from the Danian, Khazni-don River, North Osetia,
Russia: 1 – umbilical view, 2 – axial view, 3 – spiral view. L – G. planocompressa (Shutskaya), from the S. triloculinoides Subzone
(P. pseudobulloides Zone), El Kef, Tunisia: 1 – umbilical view, 2 – axial view, 3 – spiral view. M – G. planocompressa, transitional to
G. compressa, from the S. triloculinoides Subzone (P. pseudobulloides Zone), El Kef, Tunisia: 1 – umbilical view, 2 – axial view, 3 – spi-
ral view. N – G. planocompressa, transitional to G. compressa, from the E. trivialis Subzone (P. pseudobulloides Zone), Aïn Settara, Tunisia:
1 – umbilical view, 2 – axial view, 3 – spiral view.

242

ARENILLAS and ARZ

GEOLOGICA CARPATHICA

GEOLOGICA CARPATHICA, 2013, 64, 3, 237—251

Fig. 5. SEM images of Globanomalina compressa (Plummer, 1927) and Praemurica Olsson, Hemleben, Berggren & Liu, 1992 species of the
Danian (scale bar = 100 µm). A – G. compressa, holotype, from the Danian, Midway Formation, Navarro County, Texas (SEM-image from
Olsson et al. 1999): 1 – umbilical view, 2 – axial view, 3 – spiral view. B – G. compressa, from the G. compressa Subzone (P. pseudobul-
loides Zone), Ben Gurion, Israel: 1 – umbilical view, 2 – axial view, 3 – spiral view. C – Praemurica taurica (Morozova, 1961), holotype,
from the lower Danian, Tarkhankhut Peninsula, Crimea, Ukraine (SEM-image from Olsson et al. 1999): 1 – spiral view, 2 – axial view,
3 – umbilical view. D – Pr. taurica, from the Danian, Site 305, Shatsky Rise, North Pacific: 1 – spiral view, 2 – axial view, 3 – umbilical
view. E – Praemurica pseudoinconstans (Blow, 1979), holotype, from Zone P1, DSDP Leg 6, South Pacific: umbilical view. F – Pr.
pseudoinconstans, from the Danian, DSDP Site 305, Shatsky Rise, North Pacific: 1 – spiral view, 2 – axial view, 3 – umbilical view.
G – Pr. pseudoinconstans, from the S. triloculinoides Subzone (P. pseudobulloides Zone), El Kef, Tunisia: 1 – umbilical view, 2 – axial
view, 3 – spiral view. H – Praemurica inconstans (Subbotina, 1953), holotype, from Danian, Kuban River, Northern Caucasus, Russia
(SEM-image from Olsson et al. 1999): 1 – umbilical view, 2 – axial view, 3 – spiral view. I – Pr. inconstans, from the S. triloculinoides
Subzone (P. pseudobulloides Zone), Aïn Settara, Tunisia: 1 – spiral view, 2 – umbilical view. J – Pr. inconstans, from S. triloculinoides
Subzone (P. pseudobulloides Zone), El Kef, Tunisia: 1 – spiral view, 2 – axial view, 3 – umbilical view. K – Pr. inconstans, from the
E. trivialis Subzone (P. pseudobulloides Zone), Loma Capiro, Cuba: 1 – axial view, 2 – umbilical view. L – Pr. inconstans, from the
Danian, DSDP Site 305, Shatsky Rise, North Pacific: 1 – umbilical view, 2 – axial view, 3 – spiral view.

243

EARLY DANIAN NON-SPINOSE PITTED—CANCELLATE PLANKTONIC FORAMINIFERA

GEOLOGICA CARPATHICA

GEOLOGICA CARPATHICA, 2013, 64, 3, 237—251

trochospiral  species  in  the  earliest  Danian,  one  with  a
smooth wall texture (with pore-murals) and other with a rug-
ose wall texture (with rugosities and irregular pore-mounds).
The former evolved at the P0—P  transition, and was attrib-
uted to the parvularugoglobigerinids. The latter evolved at the
P —P1  transition,  and  was  assigned  to  the  new  genus  Tro-
choguembelitria Arenillas, Arz & Nán

~ez, 2012. On the con-

trary, Olsson et al. (1999) claim that the latter group belongs
to Parvularugoglobigerina, and that its wall texture is irreg-
ular pore-mounded. However, textural and biostratigraphic
data suggest that the two groups represent two different lin-
eages, Parvularugoglobigerina appearing earlier than Tro-
choguembelitria.

Evolution at the genus level

Specimens with intermediate characteristics have been re-

corded  in  El  Kef  and  Aïn  Settara  (Figs. 6—8),  illustrating
transitions  that  indicate  an  evolution  from  Parvularugoglo-
bigerina to Globanomalina, and then to Praemurica. Transi-
tional  specimens  suggest  the  following  initial  evolutionary
trends  within  this  lineage:  (1)  increase  of  test  size;  (2)  in-
crease of lip thickness; (3) increase of pore size; (4) evolution
from  tiny  pore-murals  to  large  pore-pits;  and  (5)  evolution
from smooth to cancellate wall. These evolutionary steps are
recorded in the lower part of the E. simplicissima Zone, oc-
curring quite rapidly in terms of geological time (probably in
less  than  10 kyr  according  to  the  biochronological  calibra-
tions by Arenillas et al. 2004; see Fig. 2). Within the genera,
changes in the overall shape of the test of the species seem to
be  preceded  by  the  changes  of  the  wall  textures.  Our  data
suggest an evolution from the smooth wall of Parvularugo-
globigerina to the pitted wall of Globanomalina, and finally
to the cancellate wall of Praemurica (Fig. 9).

The increase in test and pore sizes preceded the evolution of

wall texture. Several Pv. eugubina specimens show a progres-
sive  change  from  100—130 µm-length  tests  with  < 1 µm-dia-
meter  mural  pores  (Fig. 6A—B;  similar  to  the  diagram  in
Fig. 9A)  to  150—180 µm-length  tests  with  1—2 µm-diameter
mural pores (Fig. 6D; similar to diagram in Fig. 9B). The lat-
ter are clearly intermediate morphotypes between Pv. eugu-
bina  and  G.  archeocompressa,  which  still  retains  primitive
characters  such  as  mural  pores  instead  of  pore-pits  and  a
smooth instead of a pitted wall.

In the lower part of the E. simplicissima Subzone, some

Globanomalina  specimens  show  different  degrees  of  the  de-
velopment of a pitted wall (Fig. 7). Ancestral morphotypes of
G. archeocompressa (Fig. 7A) offer examples of the evolution
from mural pores of parvularugoglobigerinids to pore-pits typ-
ical of the non-spinose lineage (as in Fig. 9C). The specimen
in  Fig. 7A  still  retains  an  aperture  with  primitive  features,
namely an unusually high arch with a thin lip and therefore it
could  also  belong  to  Pv.  eugubina  or  to  Pv.  longiapertura
with  modern  textural  characters.  The  apertural  shape  of  this
specimen  is  more  common  in  the  morphotypes  intermediate
between  Pv.  longiapertura  (Fig. 3J—K)  and  Pv.  eugubina
(Fig. 3L—O) than in those between Pv. eugubina and G. ar-
cheocompressa. Blow (1979) illustrated a paratype of G. ar-

cheocompressa (Fig. 4A) with almost globular chamber and a
thick lip and considered it as a “primitive” morphotype. This
specimen seems to be an intermediate form between Pv. eugu-
bina  and  G.  archeocompressa.  Other  specimens,  like  that  in
Fig. 7B,  are  typical  examples  of  G.  archeocompressa  (with
pitted  wall,  low-arched  aperture,  thick  lip,  and  imperforated
peripheral  band  in  the  first  chambers  of  the  last  whorl),
showing different stages of development of pore-pits as ex-
emplified in Fig. 9C and 9D. The external funnel-shaped de-
pressions  of  these  pores  can  reach  up  to  about  5 µm  in
diameter (larger than in the holotype of G. archeocompressa).
These morphotypes did not develop interpore ridges, which
mean a  cancellate wall, but the degree of pore-pit develop-
ment  in  some  of  them  (e.g.  specimen  in  Fig. 7C)  strongly
supports  an  evolution  from  Globanomalina  to  Praemurica,
particularly from G. archeocompressa to Pr. taurica.

Typical Pr. taurica specimens are illustrated in Fig. 8 – the

degree  of  development  of  praemuricate  cancellate  wall  var-
ies  from  one  specimen  to  another.  Interpore  ridges  in  the
specimens  of  Fig. 8A—B  are  still  underdeveloped,  but  there
is  a  clear  tendency  to  develop  a  coarsely  cancellate  wall  as
exemplified in Fig. 9E. The overall morphology of primitive
specimens of Pr. taurica is almost identical with that of Pv.
eugubina  and  G.  archeocompressa  (Figs. 6—7).  All  three
species  have  subglobular  chambers  which  may  be  com-
pressed in some specimens.

Species of the non-spinose lineage (Globanomalina and

Praemurica) usually exhibit 5 to 6 chambers in the first whorl
(neanic stage). This character separates them from other can-
cellate Paleocene taxa such as Eoglobigerina or Parasubbotina
with  3.5  to  5  neanic  chambers  (Arenillas  &  Arz  2013).  This
difference  is  similar  to  that  which  enabled  Arenillas  et  al.
(2007) to separate  Palaeoglobigerina  (3.5  to  4  neanic  cham-
bers)  from  Parvularugoglobigerina  (4  to  4.5  chambers,  or
even 5 in the neanic stage). This morphological feature again
suggests that both Globanomalina and Praemurica are phylo-
genetically related, and that the non-spinose lineage originated
from  Parvularugoglobigerina,  with  only  a  slight  increase  in
the number of chambers in the neanic stage.

Discussion and conclusions

According to the textural, morphological and biostrati-

graphic  data,  we  propose  that  the  non-spinose  lineage  de-
rived from Parvularugoglobigerina eugubina (near the base
of  the  Eoglobigerina  simplicissima  Subzone;  Fig. 10).  Glo-
banomalina  was  the  first  taxa  of  this  lineage,  developing
pore-pits  and  acquiring  a  pitted  wall  texture.  A  short  time
later in evolutionary and geological terms (Fig. 2), cancellate
walled Praemurica evolved from Globanomalina (in the up-
per part of the E. simplicissima Subzone; Fig. 10). According
to  this  hypothesis,  truncorotalids  derived  from  globanoma-
linids,  and  these  from  parvularugoglobigerinids.  Since  their
divergence, Globanomalina and Praemurica followed sepa-
rate evolutionary paths that led to species of the two Paleo-
cene  non-spinose  lineages,  namely  the  “non-spinose  pitted”
(family Globanomalinidae) and the “non-spinose cancellate”
(family  Truncorotaloididae)  lineages.  The  oldest  species  of

247

EARLY DANIAN NON-SPINOSE PITTED—CANCELLATE PLANKTONIC FORAMINIFERA

GEOLOGICA CARPATHICA

GEOLOGICA CARPATHICA, 2013, 64, 3, 237—251

Fig. 9. Diagrams of wall textures illustrating the hypothetical evo-
lution from the smooth wall of Parvularugoglobigerina to the pitted
wall  of  Globanomalina  and  finally  to  the  cancellate  wall  of  Prae-
murica. A – smooth wall texture with tiny pore-murals (in typical
Parvularugoglobigerina).  B  –  smooth  wall  texture  with  larger
pore-murals (in more evolved Parvularugoglobigerina). C – pitted
wall texture with pore-pits (in ancestral Globanomalina). D – pit-
ted  wall  texture  with  larger  pore-pits  (in  typical  Globanomalina).
E  –  cancellate  wall  texture  (in  typical  Praemurica).  a  –  mural
pores, b – pore-pits, c – interpore ridges.

both Globanomalina and Praemurica, namely G. archeo-
compressa and Pr. taurica, initially preserved a similar test
shape of the ancestral Pv. eugubina.

Olsson et al. (1999) suggested Hedbergella holmdelensis

as  the  ancestor  of  Globanomalina  considering  that  G.  ar-
cheocompressa  exhibits  similar  features  (mainly  pitted
smooth wall and the imperforated peripheral band). However,
their  illustrated  specimens  of  G.  archeocompressa  (Fig. 3G
to  3I)  from  Zone  P0  at  Millers  Ferry  (Alabama)  do  not  fit
well  the  diagnostic  features  of  this  species  according  to  the
original description of Blow (1979). These specimens might
better  be  assigned  to  the  earliest  Danian  H.  holmdelensis,
since they have smooth—pitted more or less pustulate walls,
subconical  chambers  and  planoconvex  tests  (compare  with
H.  holmdelensis  specimens  of  Fig. 3C  to  3F).  Blow  (1979)
described  the  chamber  shape  of  G.  archeocompressa  as  al-
most  hemispherical,  the  latest  chambers  being  subglobular
(or  slightly  compressed)  with  little  or  no  differentiation  in
the  degree  of  inflation  of  their  ventral  and  dorsal  surfaces.
Arenillas  (1996)  adopted  Blow’s  concept,  illustrating  some
G.  archeocompressa  specimens  with  essentially  globular
chambers in the last whorl similar to those of Pv. eugubina.
Some  of  these  morphotypes  were  probably  considered  by
Blow  (1979)  and  Olsson  et  al.  (1999)  as  belonging  to  G.
planocompressa.

Biostratigraphical, textural and morphological data seem to

indicate that the evolutionary transition from Parvularugoglo-
bigerina to Globanomalina is more plausible than from Hed-
bergella. High-resolution biostratigraphy in Tunisia and Spain
suggested  that  the  stratigraphic  ranges  of  Hedbergella  and
Globanomalina  do  not  overlap  (Fig. 2),  except  for  some
questionable  specimens  of  Hedbergella  in  the  Pv.  eugubina
Zone,  which  were  considered  later  as  possibly  reworked
(Molina  et  al.  1996).  The  first  Globanomalina  specimens,
assigned to G. archeocompressa by Arenillas (1996), exhibit
subglobular  –  or  slightly  ovoid  –  chambers  similar  to  Pv.
eugubina.  Globanomalina  textural  features  seem  to  be  more
similar to those of Hedbergella, as proposed by Olsson et al.
(1994, 1999), except for the presence of pustules in the latter.

According to our biostratigraphic data (Fig. 2), the order

of appearance of the Globanomalina species was as follows:
G. archeocompressa (base of E. simplicissima Subzone), G.
imitata (lower part of E. simplicissima Subzone), G. plano-
compressa  (base  of E.  trivialis  Subzone)  and G.  compressa
(base of G. compressa Subzone). The evolutionary relation-
ships  between  them  are  not  well  known.  We  suggest  two
trends within early Danian Globanomalina: (1) one towards
a reduction of the number of chambers (from G. archeocom-
pressa  to  G.  imitata);  and  (2)  another  one  towards  a  com-
pressed  biconvex  shape  (from  G.  archeocompressa  to  G.
compressa).  Many  specimens  of  these  species  also  devel-
oped  an  imperforate  peripheral  band,  especially  among  the
more compressed species (G. planocompressa, G. compressa)
and also in G. archeocompressa (Fig. 7B). This feature was
considered  by  Olsson  et  al.  (1992,  1999)  to  be  directly  de-
rived from H. holmdelensis, but it could also be a character
which  reappeared  in  Globanomalina.  Olsson  et  al.  (1999)
did not place G. imitata in the lower Danian, suggesting that
it first occurred in the middle Danian (Zone P1c). Specimens

248

ARENILLAS and ARZ

GEOLOGICA CARPATHICA

GEOLOGICA CARPATHICA, 2013, 64, 3, 237—251

assignable to this species, however, have been found in older
stratigraphic levels (e.g. Fig. 4H—I), from the E. simplicissima
Subzone, or upper part of Pa, according to Arenillas et al.
(2004).

More uncertain is the evolutionary position of G. plano-

compressa,  which  seems  to  exhibit  a  morphological  feature
common  to  both  trends.  Its  author,  Shutskaya  (1965),  de-
scribed  the  chambers  of  G.  planocompressa  with  a  broadly
rounded periphery and a bluntly acute margin. Its quasi-pla-
noconvex test seems to be similar to some specimens of  G.
imitata,  suggesting  an  evolution  from  the  latter  to  the  first
one. In addition, many G. planocompressa specimens, such
as the holotype, have 4.5 chambers, suggesting that they de-
rived  from  G.  imitata  by  increasing  again  the  number  of
chambers in the last whorl. However, we suggest that G. pla-
nocompressa  is  an  intermediate  step  between  G.  archeo-
compressa  and  G.  compressa  (Fig. 10),  leading  to  an  ever
more  compressed  test,  because  some  G.  planocompressa
specimens (e.g. Fig. 4M) display shapes that already point to
G.  compressa.  Identical  phylogenetic  relationships  among

Fig. 10. Hypothetical phylogeny of the “non-spinose lineage” based on the textural and morphological characters. Phylogenetic relation-
ships of Guembelitria Cushman, 1933, Palaeoglobigerina Arenillas, Arz & Nán

~ez, 2007, Eoglobigerina Morozova, 1959 (oldest genus of

the spinose lineage) and Parvularugoglobigerina are based on Arenillas et al. (2007, 2010).

these Globanomalina species were suggested by Apellániz et
al. (2002).

Biostratigraphic data also seem to suggest that both G.

imitata  and  G.  planocompressa  derived  from  G.  archeo-
compressa, evolving in parallel to an increasingly compressed
test.  Consequently,  the  evolution  of  G.  compressa  from  G.
planocompressa  is  the  most  plausible  hypothesis.  Olsson  et
al. (1999) proposed a slightly different scenario, as they con-
sider that both G. planocompressa and G. compressa derived
independently  from  G.  archeocompressa  and  evolved  sepa-
rately.  These  hypotheses  are  not  contradictory,  since  the
taxonomic  concepts  of  G.  archeocompressa  and  G.  plano-
compressa  of  Olsson  et  al.  (1999)  differ  from  those  of
Arenillas (1996). They considered that G. archeocompressa
had  a  planoconvex  test  and  subconical  chambers  similar  to
H. holmdelensis (Fig. 3C—E), while  G. planocompressa had
subglobular to ovoid chambers, lumping morphotypes attrib-
uted  to  G.  archeocompressa  or  G.  planocompressa  by
Arenillas (1996) depending on the degree of compression of
the latest chambers (see Appendix 1).

249

EARLY DANIAN NON-SPINOSE PITTED—CANCELLATE PLANKTONIC FORAMINIFERA

GEOLOGICA CARPATHICA

GEOLOGICA CARPATHICA, 2013, 64, 3, 237—251

The evolution of the non-spinose cancellate lineage (Prae-

murica) shows an opposite trend to that of Globanomalina,
namely  the  chambers  became  more  and  more  inflated.  The
order of appearance of the Praemurica species is as follows:
Pr.  taurica  (upper  part  of  E.  simplicissima  Subzone),  Pr.
pseudoinconstans (top of E. simplicissima Subzone) and Pr.
inconstans (middle part of E. trivialis Subzone). The second
species  displays  a  feature  intermediate  between  Pr.  taurica
and Pr. inconstans. In Pr. pseudoinconstans, the last 2 or 3
chambers are strongly inflated, whereas the others are simi-
lar in size to those of Pr. taurica. In contrast, all chambers of
the  last  whorl  are  inflated  in  Pr.  inconstans.  Olsson  et  al.
(1999) suggested that the first appearance of Pr. inconstans
is  more  modern,  recorded  at  the  base  of  Zone  P1c.  In  con-
trast, we have identified abundant specimens of this species
from  the  uppermost  part  of  Zone  P1a.  This  discrepancy  is
probably  caused  by  different  concepts  of  Pr.  inconstans
among  authors  (see  Arenillas  2011).  The  separation  of  the
different species of this lineage is somewhat arbitrary, since
they  are  linked  by  transitional  morphotypes  and  their  origi-
nal  definitions  are  rather  ambiguous.  However,  these  dis-
crepancies  have  no  major  phylogenetic  implications.  All
authors agree that Pr. taurica, Pr. pseudoinconstans and Pr.
inconstans  represent  successive  evolutionary  steps  of  the
“non-spinose cancellate” lineage (Arenillas 1996; Olsson et
al. 1999; Apellániz et al. 2002).

Acknowledgments: We thank Hanspeter Luterbacher, and
an anonymous reviewer for helpful comments. We also
thank Carolina Nán

~ez for the previous review of the manu-

script.  This  research  was  funded  by  the  Spanish  Ministerio
de  Ciencia  e  Innovación  Projects  CGL2011-23077  and
CGL2011-22912 (both cofinanced by the European Regional
Development Fund), and by the Aragonian Departamento de
Educación y Ciencia (DGA group E05). The authors would
like  to  acknowledge  the  use  of  the  Servicio  General  de
Apoyo  a  la  Investigación  –  SAI,  Universidad  de  Zaragoza.
The authors are grateful to Richard Stephenson for improve-
ment of the English text.

References

Apellániz E., Orue-Etxebarría X. & Luterbacher H.P. 2002: Evolu-

tion of the early Paleocene planktonic foraminifera: a Basque
point of view. Neu. Jb. Geol. Paläont., Abh. 225, 157—194.

Arenillas I. 1996: Planktic foraminifera of the Paleocene—lower

Eocene: Systematic, biostratigraphy, chronostratigraphy and
paleoceanography. Ph.D. Thesis, Universidad de Zaragoza,
Spain, 1—513 (in Spanish).

Arenillas I. 2011: Paleoecological analysis of planktic foraminifera

of the Danian-Selandian transition from Tethys and its taxo-
nomic implications. Rev. Espan

~ola Micropaleont. 43, 109—139

(in Spanish).

Arenillas I. & Arz J.A. 1996: Origin and phylogeny of the first plank-

tic foraminiferal species of the basal Paleocene after the Creta-
ceous/Tertiary boundary. In: Perejón A. et al. (Eds.): XII Bienal,
Tomo extraordinario del 125 Aniversario de la Real Sociedad
Espan

~ola de Historia Natural, Madrid, 267—27 (in Spanish).

Arenillas I. & Arz J.A. 2000: Parvularugoglobigerina eugubina

type-sample at Ceselli (Italy): planktic foraminiferal assem-

blage and lowermost Danian biostratigraphic implications. Riv.
Ital. Paleont. Stratigr. 106, 3, 379—390.

Arenillas I. & Arz J.A. 2013: Origin and evolution of the planktic

foraminiferal Family Eoglobigerinidae Blow (1979) in the early
Danian (Paleocene). Rev. Mexicana de Ciencias Geol., 30, 1,
159—177.

Arenillas I. & Molina E. 1997: Quantitative analysis of the planktic

foraminifera from Paleocene of Caravaca (Betic Cordillera):
Biostratigraphy and faunal turnover. Rev. Espan

~ola Paleont.

12, 2, 207—232 (in Spanish).

Arenillas I., Arz J.A. & Molina E. 2004: A new high-resolution

planktic foraminiferal zonation and subzonation for the lower
Danian. Lethaia 37, 79—95.

Arenillas I., Arz J.A. & Nán

~ez C. 2007: Morphology, biometry and

taxonomy of the lowermost Danian planktonic foraminifera:
Palaeoglobigerina n. gen. Rev. Espan

~ola Paleont. 22, 21—62

(in Spanish).

Arenillas I., Arz J.A. & Nán

~ez C. 2010: Diversity and evolution of

wall textures in guembelitriids (planktic foraminifera) across
the Cretaceous-Paleogene transition. Rev. Espan

~ola Paleont.

25, 2, 87—105 (in Spanish).

Arenillas I., Arz J.A. & Nán

~ez C. 2012: Smooth and rugose wall

textures in earliest Danian trochospiral planktic foraminifera
from Tunisia. Neu. Jb. Geol. Paläont., Abh. 266, 123—142.

Berggren W.A. 1962: Some planktonic foraminifera from the Mae-

strichtian and type Danian stages of Southern Scandinavia.
Stockholm Contr. Geol. 9, 1—18.

Berggren W.A. 1977: Atlas of Paleogene planktonic foraminifera.

Some species of the genera Subbotina, Planorotalites, Morozo-
vella, Acarinina and Truncorotaloides. In: Ramsay A.T.S. (Ed.):
Oceanic micropaleontology. Academic Press, London, 250—299.

Berggren W.A. & Pearson P.N. 2005: A revised tropical to subtrop-

ical Paleogene planktonic foraminiferal zonation. J. Foram.
Res. 35, 279—298.

Blow W.H. 1979: The Cainozoic Globigerinidae. A study of the

morphology, taxonomy, evolutionary relationship and the
stratigraphical distribution of some Globigerinidae (mainly Glo-
bigerinacea). E.J. Brill, Leiden, Netherlands 3, 1—1413.

Brotzen F. & Pozaryska T. 1961: Foraminifères du Paléocène et de

l’Éocène inférieur en Pologne septentrionale; remarques
paléogéographiques. Rev. Micropaléont. 4, 155—166.

Brönnimann P. & Brown N.K. Jr. 1958: Hedbergella, a new name

for a Cretaceous planktonic foraminiferal genus. J. Washington
Acad. Sci. 48, 15—17.

Cushman J.A. 1927: An outline of a re-classification of the fora-

minifera. Contr. Cushman Laboratory Foram. Res. 3, 1—105.

Cushman J.A. 1933: Some new foraminiferal genera. Contr. Cushman

Laboratory Foram. Res. 9, 32—38.

Cushman J.A. & Bermúdez P.J. 1949: Some Cuban species of

Globorotalia. Contr. Cushman Laboratory Foram. Res. 25,
26—45.

D’Hondt S.L. 1991: Phylogenetic and stratigraphic analysis of ear-

liest Paleocene biserial and triserial planktonic foraminifera. J.
Foram. Res. 21, 168—181.

Georgescu M.D. 2007: Taxonomic re-evaluation of the Late Creta-

ceous serial planktonic foraminifer Gümbelina punctulata
Cushman, 1938 and related species. Rev. Espan

~ola Micropaleont.

39, 155—167.

Georgescu M.D. 2009a: Taxonomic revision and evolutionary clas-

sification of the biserial Cretaceous planktic foraminiferal genus
Laeviheterohelix Nederbragt, 1991. Rev. Mexicana Ciencias
Geol. 26, 315—334.

Georgescu M.D. 2009b: On the origins of Superfamily Heteroheli-

cacea Cushman, 1927 and the polyphyletic nature of planktic
foraminifera. Rev. Espan

~ola Micropaleont. 41, 107—144.

Haque A.F.M.M. 1956: The smaller Foraminifera of the Ranikot

250

ARENILLAS and ARZ

GEOLOGICA CARPATHICA

GEOLOGICA CARPATHICA, 2013, 64, 3, 237—251

and the Laki of the Nammal Gorge, Salt Range. Paleontologica
Pakistanica, Geol. Surv. Pakistan 1, 1—300.

Hofker J. 1978: Analysis of a large succession of samples through

the Upper Maastrichtian and the Lower Tertiary of Drill Hole
47.2, Shatsky Rise, Pacific, Deep Sea Drilling Project. J. Foram.
Res. 8, 46—75.

Liu C. & Olsson R.K. 1992: Evolutionary radiation of microperfo-

rate planktonic foraminifera following the K/T mass extinction
event. J. Foram. Res. 22, 328—346.

Liu C. & Olsson R.K. 1994: On the origin of Danian normal perfo-

rate planktonic foraminifera from Hedbergella. J. Foram. Res.
24, 61—74.

Loeblich A.R. Jr. & Tappan H. 1961: Suprageneric classification of

the Rhizopodea. Micropaleontology 35, 245—330.

Loeblich A.R. Jr. & Tappan H. 1984: Suprageneric classification of

the Foraminiferida (Protozoa). Micropaleontology 30, 1—70.

Loeblich A.R. Jr. & Tappan H. 1987: Foraminiferal genera and their

classification. Van Nostrand Reinhold Company, New York, 2,
1—970.

Luterbacher H.P. & Premoli Silva I. 1964: Biostratigraphy of the

Cretaceous-Tertiary boundary in Central Apennines. Riv. Ital.
Paleont. Stratigr. 70, 67—128 (in Italian).

MacLeod N. 1993: The Maastrichtian-Danian radiation of triserial

and biserial planktic foraminifera: Testing phylogenetic and
adaptational hypotheses in the (micro) fossil record. Mar. Micro-
paleont. 21, 47—100.

Molina E., Alegret L., Arenillas I., Arz J.A., Gallala N., Hardenbol J.,

von Salis K., Steurbaut E., Vandenberghe N. & Zaghbib-Turki
D. 2006: The global stratotype section and point of the Danian
Stage (Paleocene, Paleogene, “Tertiary”, Cenozoic) at El Kef,
Tunisia: original definition and revision. Episodes 29, 263—278.

Montanaro Gallitelli E. 1957: A revision of the foraminiferal Family

Heterohelicidae. Stud. Foram., United States Nat. Mus. Bull.
215, 133—154.

Morozova V.G. 1959: Stratigraphy of the Danian-Montian deposits

of the Crimea according to the foraminifera. Doklady Akad.
Nauk SSSR 124, 1113—1116 (in Russian).

Morozova V.G. 1961: Planktonic foraminifera of Danian-Montian

of the southern USSR. Paleontologicheskiy Zhurnal 1, 8—19
(in Russian).

Nederbragt A.J. 1991: Late Cretaceous biostratigraphy and develop-

ment of Heterohelicidae (planktic foraminifera). Micropaleon-
tology 37, 329—372.

Olsson R.K. 1960: Foraminifera of latest Cretaceous and earliest

Tertiary age in the New Jersey Coastal Plain. J. Paleontology
34, 1—58.

Olsson R.K. 1963: Latest Cretaceous and earliest Tertiary stratigra-

phy of New Jersey Coastal Plain. Bull. Amer. Assoc. Petrol.
Geol. 47, 643—665.

Olsson R.K. 1964: Late Cretaceous planktonic foraminifera from

New Jersey and Delaware. Micropaleontology 10, 157—188.

Olsson R.K. 1970: Planktonic foraminifera from base of Tertiary

Millers Ferry, Alabama. J. Paleontology 44, 598—604.

Olsson R.K., Hemleben C., Berggren W.A. & Liu C. 1992: Wall

texture classification of planktonic foraminifera genera in the
Lower Danian. J. Foram. Res. 22, 195—213.

Olsson R.K., Hemleben C., Berggren W.A. & Huber B.T. 1999:

Atlas of Paleocene Planktonic Foraminifera. Smithsonian Contr.
Paleobiol. 85, 1—252.

Plummer H.J. 1927: Foraminifera of the Midway formation in Texas.

Bull. Univ. Texas Bureau Econ. Geol. and Technology 2644,
1—206.

Premoli Silva I. 1977: The earliest Tertiary Globigerina eugubina

zone: Paleontological significance and geographical distribu-
tion. In: Memorias del Segundo Congreso Latinomericano de
Geología 3, Spec. Publ. 7, 1541—1555.

Sexton P.F., Wilson P.A. & Pearson P.N. 2006: Microstructural and

geochemical perspectives on planktic foraminiferal preserva-
tion: “Glassy” versus “Frosty”. Geochemistry, Geophysics,
Geosystems 7, Q12P19.

Shutskaya E.K. 1965: Phylogenetic relations of the species of the

group of Globorotalia compressa Plummer in Danian time and
the Paleocene epoch. Akad. Nauk SSSR, Voprosy Mikropaleont.
9, 173—188 (in Russian).

Smit J. 1982: Extinction and evolution of planktonic foraminifera

after a major impact at the Cretaceous/Tertiary boundary.
Geol. Soc. Amer., Spec. Pap. 190, 329—352.

Subbotina N.N. 1953: Fossil Foraminifera of the USSR. Globige-

rinidae, Hantkeninidae and Globorotalidae. Trudy Vsesoyuzhnyy
Neftyanoy Nauchno-issledovatel’skiy Geologo-razvedochnogo
Instituta (VNIGRI), Mikrofauna SSSR Sbornik 6, 76, 1—296 (in
Russian).

251

EARLY DANIAN NON-SPINOSE PITTED—CANCELLATE PLANKTONIC FORAMINIFERA

GEOLOGICA CARPATHICA

GEOLOGICA CARPATHICA, 2013, 64, 3, 237—251

Diagnostic characters of the early Danian species of Hedbergella Brönnimann & Brown, 1958, Parvularugoglobigerina

Hofker, 1978, Globanomalina Haque, 1956 and Praemurica Olsson, Berggren & Liu, 1992 mentioned in the text (Arenillas
1996, modified):

Appendix

Hedbergella monmouthensis Olsson (1960): Trochospiral

test, 5—6 hemispherical to subconical chambers in the first
whorl, 4—5 subglobular chambers in the last whorl, moderate
rate of chamber size increase, axial periphery rounded, aper-
ture umbilical—extraumbilical, with thick lip, wall surface
smooth, pustulate (Fig. 3A—B).

Hedbergella holmdelensis Olsson (1964): Trochospiral test,

5—6 hemispherical chambers in the first whorl, 5—6.5 moder-
ately compressed hemispherical to subconical chambers in the
last whorl, low rate of chamber size increase, axial periphery
slightly angular, aperture umbilical—extraumbilical, with thick
lip, wall surface smooth, pustulate (Fig. 3C—I).

Parvularugoglobigerina longiapertura (Blow 1978):

Small trochospiral test, with low spire, 4—4.5 moderately
compressed ovate to subglobular in the first whorl, 5—6.5
slightly to moderately compressed ovate chambers in the last
whorl, low rate of chamber size increase, aperture extending
up into the apertural chamber face, high arch, with lip thin,
wall surface smooth or secondarily granular (Fig. 3J—K).

Parvularugoglobigerina eugubina (Luterbacher & Premoli

Silva 1964): Small trochospiral test, with low spire, 4—4.5 sub-
globular chambers in the first whorl, 5—6.5 subglobular cham-
bers in the last whorl, low rate of chamber size increase,
aperture, umbilical—extraumbilical, low arch, with lip thin, wall
surface smooth or secondarily granular (Figs. 3L—O, 6A—D).

Globanomalina archeocompressa (Blow 1979): Trocho-

spiral test, with low spire, 4—4.5 subglobular chambers in the
first whorl, 5.5—6.5 subglobular to slightly ovoid chambers
in the last whorl, low rate of chamber size increase, aperture
umbilical—extraumbilical, low arch, with lip moderately
thick, wall surface pitted (Figs. 4A—F, 7A—C).

Globanomalina imitata (Subbotina 1953): Trochospiral

test, with low spire, 4—4.5 subglobular to hemispherical
chambers in the first whorl, 4 subglobular to hemispherical

to planoconvex chambers in the last whorl, moderate rate of
chamber size increase, aperture umbilical—extraumbilical,
with lip moderately thick, wall surface pitted (Fig. 4G—J).

Globanomalina planocompressa (Shutskaya 1965): Tro-

chospiral test, with flat spire, 4—4.5 hemispherical chambers
in the first whorl, 4.5—5 ovoid to hemispherical chambers in
the last whorl, low to moderate rate of chamber size increase,
aperture umbilical—extraumbilical, with lip thick, wall sur-
face pitted (Fig. 4K—N).

Globanomalina compressa (Plummer 1927): Trochospiral

test,  biconvex,  5—6  moderately  compressed  ovate  chambers
in the first whorl, 4.5—5.5 slightly to moderately compressed
ovate  chambers  in  the  last  whorl,  low  to  moderate  rate  of
chamber  size  increase,  axial  periphery  rounded  to  slightly
angular, with imperforate margin with poor developed or ab-
sent,  aperture  umbilical—extraumbilical,  with  lip  thick,  wall
surface pitted to smooth (Fig. 5A—B).

Praemurica taurica (Morozova 1961): Trochospiral test,

with low to flat spire, 5—6 subglobular chambers in the first
whorl, 5.5—7 subglobular chambers in the last whorl, low rate
of chamber size increase, aperture umbilical—extraumbilical,
with lip thick, wall surface cancellate (Figs. 5C—D, 8A—C).

Praemurica pseudoinconstans (Blow 1979): Trochospiral

test,  with  low  to  flat  spire,  5—6  subglobular  chambers  in  the
first whorl, 5—5.5 subglobular chambers in the last whorl (2 or
3  last  chambers  very  inflated),  moderate  to  high  rate  of
chamber  size  increase,  aperture  umbilical—extraumbilical,
with lip thick, wall surface cancellate (Fig. 5E—G).

Praemurica inconstans (Subbotina 1953): Trochospiral

test, with low to flat spire, 5—6 subglobular chambers in the
first whorl, 5—6.5 inflated subglobular chambers in the last
whorl, low to moderate rate of chamber size increase, aper-
ture umbilical—extraumbilical, with lip thick, wall surface
cancellate (Fig. 5H—L).