423

ATLAS SP. # 102

Duquepsammina cubensis (Cushman & Bermúdez, 1937)

Fig. 102. 1. holotype of of Spiroplectoides cubensis from Cushman & Bermúdez (1937), 2,3. microsphaeric
specimens from Seiglie & Baker (1987).

ORIGINAL DESIGNATION: Spiroplectoides cubensis Cushman & Bermúdez, 1937.

TYPE REFERENCE: Cushman, J.A. & Bermúdez, P.J. 1937. Further new species of foraminifera from the Eocene of Cuba.
Contr. Cushman Lab. Foram. Res., 13, p. 13, pl. 1, figs. 44-45. see also: Seiglie, G.A. & Baker, M.B. 1987. Duquepsamminiidae, a
new family, and Duquepsammia, a new genus of agglutinated foraminifers. Micropaleontology, 33, 263-266.

TYPE SPECIMEN: Deposited in the Cushman Collection, U.S. Natural History Museum, Smithsonian
Institution (Holotype CC 23384; Paratype 23385). Several metatype specimens are also preserved in the
collection of "secondary types", including specimens from Columbia studied by Seiglie & Baker (1987).
Eight slides of additional topotype and metatype specimens are housed in the Bermúdez Collection at
INTEVEP S.A. (Centro de Investigación y Apoyo Tecnológico, Filial de Petróleos de Venezuela, S.A) in
Caracas.

TYPE LEVEL: Late Eocene.

TYPE LOCALITY: Bermúdez Station #18, Alturas de Almendares Quarry in Reparto Kohly, west side of
the Almendares River, Havana Province, Cuba.

DIAGNOSTIC FEATURES: Test very elongate, slender, laterally compressed, with acute periphery. The
initial portion consists of a biconcave evolute planispire of as many as 3 1/2 whorls with up to 12 chambers
in the outer whorl, followed by a biserial part consisting of as many as 22 pairs of chambers. Chambers
increase in size slowly, are low in the early biserial portion, and become higher with ontogeny. The biserial
chambers are partially subdivided by constrictions formed by the thickened septal wall in the middle part of
the chamber. Sutures are depressed, distinct when viewed in immersion, becoming more limbate towards
the median axis of the biserial part, and become more oblique with ontogeny. Wall finely agglutinated with
a smooth surface. Aperture an interiomarginal slit within a slight depression; in specimens with one or two
uniserial chambers a terminal oval opening.

SIZE: Holotype is 1.54 mm in length, and 0.20 mm in maximum width, and 0.11 mm in thickness. Width
of the spiral part is 0.18 mm, and the width of the initial portion of the biserial part is 0.12 mm.
 424

SYNONYMS: None verified.

OBSERVED OCCURRENCES: Cushman & Bermúdez (1937) originally described this species as
Spiroplectoides cubensis from the Upper Eocene of Havana Province, Cuba. Additional specimens in the
Bermúdez Collection are from the Eocene and Oligocene of the Pinar de Rio Province, mostly near Mariel,
Cuba. Cushman & Stainforth (1945) subsequently reported it from the Cipero Formation of Trinidad "in all
three zones", i.e. from the Oligocene to Middle Miocene. Cushman & Renz (1947) recorded it from the
upper Oligocene St. Croix Formation of Trinidad, and in their subsequent publication (Cushman & Renz,
1948) recorded it in the Eocene Fitt Trace, Nariva River, Friendship Quarry, and Dunmore Hill marls of
Trinidad.
In the Pacific region, this species was reported as Bolivinopsis cubensis by Finlay (1946) from the
Lower Oligocene (Whaingaroan) Oxford Chalk of New Zealand, by Hornibrook (1961) from the Late
Eocene to Early Miocene of the Oamaru District, New Zealand, and by Gibson (1967) from the Upper
Miocene (Tongaporutuan) of New Zealand. Todd (1966) illustrated an atypical specimen from the
Oligocene of Guam, Thomas (1985) recorded it from the upper Eocene to lowermost middle Miocene at
DSDP Sites 573, 574, and 575 in the central equatorial Pacific, Boersma (1985) recorded it from the
uppermost Eocene to Lower Oligocene at Sites 592 and 593 in the Tasman Sea and Chen et al. (1997)
figured a specimen of early Miocene age from a piston core recovered from the CC area (between the
Clarion and Clipperton Fracture Zones) in the Northeast Pacific Basin. Kuhnt et al. (2002) illustrated
specimens at Bolivinopsis praelonga (Schwager) from the Oligocene of ODP Site 1148 in the South China
Sea.
In the Atlantic region, Miller & Katz (1987) recorded it as common in the Oligocene at DSDP Site 558,
and as rare in the Lower Oligocene to Lower Miocene at DSDP Site 563. We observed this species in the
mid Oligocene of the Cipero Formation of Trinidad. We have found rather slender specimens in the Upper
Eocene and Lower Oligocene at ODP Site 647 in the Labrador Sea.

KNOWN STRATIGRAPHIC RANGE: Late Eocene to Late Miocene (Seiglie & Baker, 1987).

BATHYMETRY: Middle Bathyal to Abyssal.

REMARKS: Duquepsammina cubensis is easily distinguished from other species by its large, evolute,
spiral part and its long, slender biserial part with numerous chambers and thickened sutures. Metatypes of
"Spiroplectoides cubensis" preserved in the Cushman Collection are from the Eocene Navet and Oligocene
Cipero Formations of Trinidad. All forms preserved in the collection are megalosphaeric.

Fig. 102-2. Drawing of the biserial stage of a specimen of D.

cubensis (USNM 401088) showing inner structure of sutures,
from Seiglie & Baker (1987).

The specimen illustrated by Cushman & Stainforth (1945) (CC43583) has 12 chambers in its spiral part, 18
pairs of chambers in its biserial part, and measures 1.43 mm. In large individuals, the sutures in the biserial
part become progressively more limbate with ontogeny. Near the median axis of the test, the sutures appear
 425
Plate 102 - Spiroplectammina cubensis (Cushman & Bermúdez)

Figs 1–2. Eocene, Habana Province, Cuba, Metatype specimens, Bermúdez Collection at INTEVEP,
Caracas; courtesy of Humberto Carvajal-Chitty. Figs. 3a,b. Oligocene, Central North Atlantic Site 558,
sample 558-26-1, 93-97 cm, courtesy of Mimi Katz, 3b. x84. Figs. 4–5. Late Eocene, Labrador Sea Site
647, sample 647A-31-1, 132-136 cm, 4. x66, 5. x100. 6a,b. Oligocene, lower part of Cipero Formation,
Trinidad, CC43586, transmitted and reflected light in immersion, x60. Figs. 7a–8b. Oligocene, upper part
of the Cipero Formation, Trinidad, CC43586, transmitted and reflected light in immersion, 7. x60. 8. x50.
 426

as thickened nodes when viewed in immersion (Fig. 102-2), partially subdividing the chambers. Based on
this criterion, Seiglie & Baker (1987) described the genus Duquepsammina, with S. cubensis as the type
species.

ILLUSTRATIONS: Plate 102 - Duquepsammina cubensis (Cushman & Bermúdez) –

ATLAS SP #103
Spiroplectammina navarroana Cushman, 1932 emend. Gradstein & Kaminski, 1989

Fig. 103-1. 1a,b. Holotype of Textularia plummerae (CC

21922); 2a,b. Holotype of Spiroplectammina navarroana.
3a,b. Paratype of S. navarroana. All figures are drawn
from the original specimens housed in the Cushman
Collection.

ORIGINAL DESIGNATION: Spiroplectammina navarroana Cushman, 1932.

TYPE REFERENCE: Cushman, J.A. 1932. Textularia and related forms from the Cretaceous. Contr. Cushman Lab. Foram.
Res., 8, p. 96, pl. 11, fig. 14. see also: Gradstein, F.M. & Kaminski, M.A. 1989. Taxonomy and biostratigraphy of new and
emended species of Cenozoic deep-water agglutinated foraminifera from the Labrador and North Seas. Micropaleontology, 35, p.
83, pl. 9, figs. 1a-12.

TYPE SPECIMEN: The holotype (USNM 371545) and paratype (USNM 26887) are housed in the
Cushman Collection, U.S. Natural History Museum, Smithsonian Institution, Washington, D.C.

TYPE LEVEL: Maastrichtian, upper clay member of the Navarro Formation.

TYPE LOCALITY: Six miles east of Corsicana, Navarro County, Texas.

DIAGNOSTIC FEATURES: Test elongate, arched, tapered, initially planispiral, later biserial, with broadly
rounded periphery. Chambers are approximately as high as they are broad and increase slowly in size as
added. The final one or two chambers may be somewhat more inflated than earlier chambers. Sutures are
distinct, initially inclined with respect to the long axis of the test, later perpendicular to the axis, becoming
more incised towards the distal end of the test. Wall imperforate, medium-coarse to finely agglutinated.
Aperture varies from a low interiomarginal arch to a higher arch.

SIZE: Mean length is 850 microns.

SYNONYMS:
Textularia plummerae Lalicker. Lalicker, C.G., 1935. Contr. Cushman Lab. Foram. Res. 11, p. 67, pl. 3,
figs. 2a,b [Eocene, Texas].
Spiroplectammina lanceolata Huss. Huss, J., 1966. Prace Geologiczne PAN, 34, p. 36, pl. 5, figs. 16-20
[Campanian, Polish Carpathians].

OBSERVED OCCURRENCES: Spiroplectammina navarroana is a ubiquitous component of flysch-type

agglutinated assemblages and occurs at nearly every locality studied. The syntypes were originally
described from the upper clay member of the Navarro Formation of Texas. Additional metatype specimens
 427

in the Cushman Collection labelled S. navarroana are from the Marlbrook Marl of Arkansas (CC42033 -
CC42035). As Textularia plummerae, it was described from the Eocene Mexia clay member of the Wills
Point Formation of Texas by Lalicker (1935). It has also been reported under this name from the Paleocene
of the Azovo-Kubansk Basin in the Don-Bass region of Russia (Nikitina & Shvemberger, 1963); from the
Paleocene (P1d) to Middle Eocene (P12) of the Possagno section in Italy (Braga et al., 1975); from the
Danian of the Buntmergelserie in Bavaria by De Klasz & De Klasz (1990), from the Paleocene to Middle
Eocene of New Zealand by Kaiho et al. (1993); from the Paleocene Jamburo Group of the Khuzdar District
in Pakistan by Brohi (1994); and from the Paleocene Moogli Mudstone of Papua New Guinea by Milner
(1997). Cushman & Renz (1946) reported it as "Gaudryina foeda" from the Lizard Springs Formation of
Trinidad. Geroch & Nowak (1984) reported this species as Spiroplectammina lanceolata in the Polish
Carpathians, where its range was reported as Turonian to Maastrichtian. Bąk (2000) reported it as S.
navarroana from Upper Cretaceous scaglia-type facies in the Pieniny Klippen Belt, and Bąk (2004)
reported it from organic-rich facies of the Paleocene Majdan Beds in the Dukla Unit.
We have observed this species in nearly every studied locality that contains a flysch-type assemblage.
In Trinidad, it occurs from the Late Maastrichtian A. mayaroensis Zone to the middle Eocene (P11) of the
Navet Formation. In the flysch deposits at Zumaya in northern Spain, it ranges from Zones P1c to P6b,
Paleocene to P/E transition. In the Central North Sea and the Labrador Margin, its average last occurrence
is in strata of late Early Eocene to early Middle Eocene age. In ODP Hole 647 in the southern Labrador
Sea, it was found in the Lower and Middle Eocene. We have observed it in the Maastrichtian to Paleocene
at ODP Site 959 in the South Atlantic, which contains a bathyal assemblage. Spiroplectammina navarroana
has not been reported from deep abyssal agglutinated assemblages described by Krasheninnikov (1973,
1974), Hemleben & Tröster (1984) and Moullade et al. (1988); and we have not observed it in our samples
from the Paleogene Scaglia Rossa at Gubbio.

KNOWN STRATIGRAPHIC RANGE: Turonian - Eocene.

BATHYMETRIC RANGE: Bathyal to upper abyssal.

REMARKS: We have formally emended this species based upon a restudy of the type material of
Cushman, Lalicker, and Huss (Gradstein & Kaminski, 1989). Cushman (1932) based his initial description
of S. navarroana on broken specimens, and made no mention of an initial planispire. Only two of
Cushman's specimens are preserved in the Cushman Collection. Cushman originally described S.
navarroana as being "very slightly if at all tapering in the adult portion". Examination of the holotype and
paratype specimen in transmitted light (textfigures 2a,b and 3a,b) reveals that they are short (0.62 and 0.47
mm) biserial fragments that are not representative of the species. Specimens from other localities are
usually longer and contain more pairs of chambers in the biserial part. Unbroken specimens are rare, but
complete specimens have a minute initial whorl of about 5 chambers and have a tapered, arching test (plate
103, fig. 4). The holotype of Textularia plummerae Lalicker (1935) from the Eocene Mexia Clay of Texas
(Fig. 103-1, 1a,b) is, in our opinion, indistinguishable from S. navarroana from North Atlantic and Tethyan
localities. The species S. lanceolata Huss (1966) was described as having 10 to 13 rows of biserial
chambers and a small initial planispiral whorl situated eccentric to the long axis of the test. The type
specimens of S. lanceolata are missing from the collections of the Polish state petroleum exploration
company "NAFTA", but our topotype specimens of S. lanceolata from the Upper Cretaceous Węglówka
Marls of Poland are identical to S. navarroana from the North Atlantic flysch-type assemblages.
Spiroplectammina navarroana is variable in the size and nature of the agglutinated grains used in the
construction of the test wall. The type specimens are comprised of relatively coarse material which includes
some mafic grains, whereas specimens from Site 647 (plate 103, fig. 3) are comprised of fine grained
pelagic material.

ILLUSTRATIONS: Plate 103 - Spiroplectammina navarroana Cushman, emend. Gradstein & Kaminski –
 428
Plate 103 - Spiroplectammina navarroana Cushman, emend. Gradstein & Kaminski

Figs. 1a–2. Paleogene, Dominion O-23 well; Fig. 3. Lower Eocene, Labrador Sea Site 647, Sample 647A-
67R-1, 40-43 cm. x140; Figs. 4–9. Campanian, W ęglówka Marls, Subsilesian Unit of the Polish
Carpathians, approx x75; Figs. 10–13. Paleocene, Lizard Springs Formation, Trinidad, x75.
 429

ATLAS SP #104
Spiroplectammina spectabilis (Grzybowski, 1898) emend. Kaminski, 1984

Fig. 104-1. Five species of Spiroplecta described by Grzybowski (1898, 1901) here regarded as synonymous with S.
spectabilis (redrawn from the type figures). 1. Spiroplecta brevis Grzybowski, 1898. 2. Spiroplecta spectabilis Grzybowski,
1898. Spiroplecta costidorsata Grzybowski, 1898. 4. Spiroplecta foliacea Grzybowski, 1898. 5. Spiroplecta clotho
Grzybowski, 1901

ORIGINAL DESIGNATION: Spiroplecta spectabilis Grzybowski, 1898.

TYPE REFERENCE: Grzybowski, J. 1898. Otwornice pokł adow naftonoś nych okolicy Krosna. Rozprawy Wydziału
Matematyczno-Przyrodniczego, Akademia Umiejętności w Krakowie, serya 2, vol. 33, p. 293, pl. 12, fig. 12. See also: Kaminski,
M.A. 1984. Shape variation in Spiroplectammina spectabilis (Grzybowski). Acta Paleontologica Polonica, 29, 29-49.

TYPE SPECIMEN: Not originally designated. Numerous specimens from Grzybowski's samples are
deposited in the micropaleontological collections of the Jagiellonian University, Kraków. Syntypes of
Spiroplecta spectabilis preserved in the Grzybowski Collection of 1898 are registered as UJ-132-P, 1/82a.
This slide contains three specimens, one of which corresponds to the type figure. Kaminski & Geroch
(1993) designated this specimen the lectotype. An additional syntype of S. spectabilis labelled "Potok" is
registered in slide UJ-132-P, 1/82b. Syntypes of Spiroplecta brevis are registered as UJ-132-P, 1/83a-e.
Syntypes of Spiroplecta foliacea are registered as UJ-132-P, 1/84a. The type specimen of Spiroplecta
costidorsata is registered as UJ-132-P, 1/85.

TYPE LEVEL: Paleogene, Silesian Unit of the Polish Carpathians.

TYPE LOCALITY: Not originally designated. Grzybowski reported Spiroplecta spectabilis from three
exploration wells drilled in the vicinity of Krosno Poland. The lectotype is from the Potok H-33 well, 170
m. Syntypes of Spiroplecta brevis are from the Potok H-33 well, 170 m, 210 m, 227 m, and 15 m.
Spiroplecta foliacea were described from an additional six wells. Syntypes in the collection are from the
Potok H-40 well, 24 m, and from an outcrop sample labelled " Krościenko, red clay". The type specimen of
Spiroplecta costidorsata is from the Potok H-26 well, 95 m.

DIAGNOSTIC FEATURES: Test free, initially planispiral, later biserial. In megalospheric forms, the
planispiral portion consists of 4-7 chambers, and may be wider than the subsequent biserial portion. The
planispiral portion of microspheric forms is minute in comparison, and may also be wider than the biserial
part. Chambers in the biserial part are low and numerous. As many as 36 biserial chambers have been
observed in microspheric individuals. In both generations, the biserial part is rhomboidal in cross-section
and has nearly parallel sides, though in microspheric forms it normally increases in breadth initially and
may decrease in breadth distally. Sutures are normally flush or depressed slightly. In the biserial part,
sutures and inclined approximately 60° to the long axis of the test. Peripheral margin is acute, and may be
dentate or weakly keeled. Wall is imperforate, finely agglutinated with a smooth surface. Forms living on
pelagic substrates may agglutinate coccolith debris. Aperture is a narrow interiomarginal slit.

SIZE: Mean length varies from 0.35 to 1.25 mm. The lectotype of S. spectabilis is 0.20 mm x 0.55 mm.
 430

SYNONYMS:
Spiroplecta brevis Grzybowski. Grzybowski, J., 1898, Rozpr. Wydz. Matemat.-Przyr., Akad. Umiej.
Krakow, ser. 2, 33, p. 293, pl. 12, fig. 13.
Spiroplecta foliacea Grzybowski. Grzybowski, J., 1898, Rozpr. Wydz. Matemat.-Przyr., Akad. Umiej.
Krakow, ser. 2, 33, p. 293, pl. 12, fig. 12.
Spiroplecta costidorsata Grzybowski. Grzybowski, J., 1898, Rozpr. Wydz. Matemat.-Przyr., Akad. Umiej.
Krakow, ser. 2, 33, p. 294, pl. 12, fig. 11.
Spiroplecta clotho Grzybowski. Grzybowski, J., 1801, Rozpr. Wydz. Matemat.-Przyr., Akad. Umiej.
Krakow, ser. 2, 41, p. 283, pl. 7, fig. 18.
Spiroplectoides clotho (Grzybowski). Cushman, J.A., & Jarvis, P.W., 1928, Contr. Cush. Lab. Foram. Res.
4, p. 101, pl. 14, fig. 13.
?Spiroplectoides eocenica Cushman & Barksdale. Cushman, J.A. & Barksdale, J.D., 1930, Contr. Stanford
Univ. Geol. Dept. 1, p. 66, pl. 12, fig. 5a,b.
Spiroplectoides californica Cushman & Campbell. Cushman, J.A., & Campbell, A.S., 1934, Contr. Cush.
Lab. Foram. Res. 10, p. 70, pl. 9, fig. 13-14.
Spiroplectammina mexiaensis Lalicker. Lalicker, C.G., 1935. Contr. Cush. Lab. Foram. Res. 11, p. 43, pl.
6, fig. 5-6.
Spiroplectoides directa Cushman & Siegfus. Cushman, J.A., & Siegfus, S.S., 1939. Contr. Cush. Lab.
Foram. Res. 15, p. 26, pl. 6, fig. 7-8.
Spiroplectammina grzybowskii Frizzell. Frizzell, D.L., 1943. J. Paleontol. 17, p. 339, pl. 55, fig. 12a-13.
Spiroplectammina perplexa Israelsky. Israelsky, M.C., 1951. U.S. Geol. Surv. Prof. Paper 240-A. p. 12, pl.
3, fig. 9-14.
Spiroplectammina brunswickensis Todd & Kniker. Todd, R., & Kniker, H.T., 1952. Cush. Found. Foram.
Res. Spec. Publ. 1, p. 6, pl. 1, fig. 16.

OBSERVED OCCURRENCES: Spiroplectammina spectabilis is one of the best known and widely
distributed Paleogene species of deep-water agglutinated foraminifera. It was first described from the
Paleogene of the Carpathian flysch (Grzybowski, 1898), and has been subsequently found in the Caucasus
(Glaessner, 1937; Subbotina, 1950), Caribbean (Cushman & Jarvis, 1928), Tunisia (Aubert & Berggren,
1976), along the Pacific margin of North and South America (Cushman & Campbell, 1934; Todd & Kniker,
1952), in the Northwest Pacific (Serova, 1987; Kaiho, 1992), South Pacific and Papua New Guinea (Webb,
1975; Haig, 1982; Milner, 1997), New Zealand (Kaiho et al., 1996), South Atlantic (Widmark, 1997), and
throughout the North Atlantic and North Sea provinces. In its type area, the first occurrence of this species
marks the base of the Late Paleocene Spiroplectammina spectabilis Zone of Geroch & Nowak (1984).
However, this event in the Polish Carpathians may reflect the immigration of the species, since it
undoubtedly occurs in the Maastrichtian in Trinidad (Kaminski et al., 1988; Beckmann, 1994), in northeast
Venezuela (Galea-Alvarez, 1985), on Sakhalin Island (Serova, 1987), and on Hokkaido (Kaiho, 1992). It is
found in the Lower Paleocene of other areas of the Carpathians as well as in the Atlantic. Martin (1964)
reported it as Spiroplectammina perplexa from the Lower Danian of the Moreno Formation in California. In
the Romanian Eastern Carpathians, Bratu (1975) noted its frequent occurrence in the upper Eocene
Bisericani Formation, and described “an assocation with S. spectabilis and Globigerinatheka index”. Ion
(1995) recorded its FO in the G. eugubina Zone in the Lepsa Beds, and noted reports of it in the uppermost
Maastrichtian of the Hangu Beds. Based on these occurrences, Ion (1995) argued for restricting the S.
spectabilis Zone in the Romanian Carpathians to the interval between the LO of Rzehakina inclusa and the
FO of Saccamminoides carpathicus.
At Caravaca Spain, the FO of S. spectabilis was observed within the K/T boundary clay layer (Coccioni
& Galeotti, 1994). We have also observed a peak in S. spectabilis above the K/T boundary in deep-water
limestones from Gubbio and from Sopelana Spain (Kuhnt & Kaminski, 1993; 1996). Peryt et al. (1997)
observed the FO of S. spectabilis in Zone PO in flysch sediments of the Rotwandgraben section, Eastern
Alps (Austria). At this locality its acme is in Zone Pα to P1a. At ODP Site 959 in the equatorial Atlantic, its
the FO of S. spectabilis is only centimeters above the K/T boundary, and it displays a prominent acme in
the lower Danian (Fig. 104-2). We observed a similar peak in the Danian in one or two Labrador Shelf
wells. This peak in the abundance of S. spectabilis following the K/T boundary event therefore appears to
have a wide geographical distribution.
 431

Depth % Spiroplectammina % Rzehakina % Caudammina

(m bsf) spectabilis epigona gigantea
0 10 20 30 0 5 10 15 0 5 10 15 20
860

Spi ropl ectammi na Event

Caudammi na ex gr.
870 gigantea Exti ncti on

880

Fig. 104-2. Abundance variations of common DWAF across the K/T boundary interval at ODP Site 959 off the Ivory Coast. The
early Danian acme in Spiroplectammina spectabilis has now been observed at several localities in the Atlantic and western Tethys,
after Kuhnt et al. (1998).

In the Zumaya section of northern Spain, its first occurrence is near the base of Zone P4. In the northern
areas of the North Atlantic, S. spectabilis can be abundant in sediments of Late Paleocene age. It displays a
distinctive abundance maximum in the lower part of Upper Paleocene in the western Barents Sea. Its last
common occurrence occurs near the base of the uppermost Paleocene tuff horizon in the North Sea and
Barents Sea, and near the Paleocene/Eocene boundary along the Labrador Margin. Another acme in S.
spectabilis has been found in the Contessa Road section near Gubbio within Zone NP9, just beneath the
Paleocene/Eocene boundary carbon isotope excursion (Galeotti et al., 2000). The last occurrence of S.
spectablis is generally within the Middle Eocene along the North Atlantic margins. However, at
paleobathymetrically deeper localities it ranges into Upper Eocene. At ODP Site 647 in the Labrador Sea its
LO coincides with the Eocene/Oligocene boundary as determined by nannofossils and magnetostratigraphy.
Verdenius & Van Hinte (1983) used this species to determine the top of the Eocene in DSDP sites in the
Norwegian-Greenland Sea. In a study of a paleobathymetric transect in the Central North Sea, Jones (1988)
reported it as common in upper slope and basin plain assemblages but less common in the middle slope
facies. In the Polish Carpathians its LO coincides with the base of the uppermost Eocene "Globigerina
Marls".
Spiroplectammina spectabilis was largely restricted to flysch-type assemblages. It has not been found in
DSDP/ODP assemblages from abyssal sites in the North Atlantic, except at high latitudes (Site 647), where
it forms a distinctive acme in Zone NP17 (see Stratigraphy chapter). Its occurrence is rare and
discontinuous in the Paleogene Scaglia Rossa of Italy, except at particular intervals. Its distribution suggests
seems to be linked with the trophic continuum. It is extremely rare in the eastern Atlantic ODP sites (Kuhnt
& Collins, 1996; Kuhnt & Urquhart, 2001). In the western Barents Sea, its maximum occurrence is
observed in the middle part of the Paleocene, while in the Central North Sea it is most common in the upper
Paleocene. Its occurrence in the deep-sea sites is most likely linked to elevated organic input and
siliciclastic flux.

KNOWN STRATIGRAPHIC RANGE: Maastrichtian to latest Eocene.

BATHYMETRY: Bathyal to abyssal.

 432

REMARKS: Spiroplectammina spectabilis may be one of the most widely recognised Paleogene species,
since it figures prominently in a number of zonal schemes. This species may be considered as a plexus of
forms that differ in length and thickness, with significant differences between end members (Kaminski,
1984). This variability has no doubt contributed to the multiplicity of available junior synonyms. The
concept of Spiroplectammina spectabilis adopted here consists of no fewer than 11 validly described
species.
Grzybowski described five species of Spiroplecta that we consider to be fully synonymous with S.
spectabilis. Spiroplecta brevis and S. spectabilis are clearly megalospheric forms, while S. costidorsata, S.
foliacea and S. clotho are microspheric individuals.
Four complete specimens of Spiroplecta brevis are preserved in the Grzybowski collections. In all
cases, Grzybowski selected specimens that are comparatively short (4-5 pairs of biserial chambers) and
possess an initial spire that is wider than the biserial part. It appears that Grzybowski himself may have had
difficulty distinguishing S. spectabilis from S. brevis, since there are undifferentiated vials in his collection
that are labelled as containing both forms.
Two specimens of Spiroplecta foliacea are preserved in the Grzybowski collections, but the complete
specimen depicted in Grzybowski's pl. 12, fig. 12 appears to be missing. Both specimens appear to be wider
than the one depicted in Grzybowski's type figure. The longest specimen is broken, and has 15 pairs of
chambers. In our understanding, the type figure of S. foliacea depicts a rather narrow microspheric
individual of Spiroplectammina spectabilis, whereas S. clotho is an abnormally wide specimen. The
surviving specimen of Spiroplecta costidorsata preserved in the Grzybowski collection is a microsphaeric
individual with 13 pairs of chambers. Unlike the type figure, the preserved syntype has parallel sides
without any trace of lateral spines, as the name and the type illustration suggests.
In their studies of faunas from Trinidad, Mexico, and California, Cushman and co-workers used the
designations Spiroplectoides clotho (Grzybowski) and S. spectabilis, but described three additional species
from California: S. eocenica, S. californica and S. directa (Fig. 104-3). A fourth species from California
was described by Israelsky (1951).
Spiroplectoides eocenica was described by Cushman & Barksdale (1930) from the Eocene Martinez
Formation of the San Francisco area. Cushman & Barksdale reported that it "resembles Spiroplectoides
clotho (Grzybowski), but the chambers are lower, the sides parallel, the sutures hardly if at all limbate, and
the relative length of the test is very much less". The wall was described as being "distinctly perforate". We
have discovered, however, that Cushman often mistook the fine agglutinated grains for perforations.
Therefore we tentatively include this species as a synonym of S. spectabilis pending revision of the types.
Spiroplectoides californica was described by Cushman & Campbell (1934) from the Paleogene Chico
and Moreno Formations. The species was described as "varying in thickness at the apertural end, which
becomes as thick as broad". In comparison to specimens from Trinidad that Cushman had placed in S.
clotho, the specimens from California were described as having chambers that are "not so low and broad as
in the latter species, and the microspheric form is not so definitely lance-shaped". However, the type
specimens of S. californica preserved in the Cushman Collection (CC21294 - CC21299) are not as thick as
they are broad. The holotype is 0.28 mm in width and only 0.21 mm in thickness. The mean thickness of
the five preserved type specimens is 0.91 ± 0.2 mm. The mean thickness of these specimens is not
significantly different from virtual topotypes of S. spectabilis collected from the Polish Carpathians.
Spiroplectoides directa was described by Cushman & Siegfus (1939) from the Middle to Upper Eocene
Kreyenhagen Shale at Garza Creek, California. The species was described as having a thin, perforate,
calcareous wall and a terminal aperture. The holotype (CC 25445) is apparently silicified with an
agglutinated test, and the specimen is broken at its apertural end. This gives it the appearance of having a
terminal aperture. The species was again reported by Cushman & Siegfus (1942) from the Kreyenhagen
Shale type locality. Cushman & Renz (1948) later used this name for specimens from the Middle Eocene
Navet Formation of Trinidad.
 433

Fig. 104-3. Specimens of S. spectabilis from localities in the Americas illustrated by Cushman (1927, 1934), Cushman & Campbell
(1934) and Cushman & Siegfus (1939), redrawn from Cushman's illustrations. 1-5. Specimens reported as "Spiroplectoides clotho".
Specimen #3 is from the Velasco Shale of Mexico, others are from Trinidad. 6-7. Specimens reported as "Spiroplectoides
spectabilis" by Cushman (1934). 8. Specimen reported as "Spiroplectoides eocenica" by Cushman & Barksdale (1930). 9-10.
Specimens reported as "Spiroplectoides californica" by Cushman & Campbell (1934). 11-12. Specimens reported as
"Spiroplectoides directa" by Cushman & Siegfus (1939).

Spiroplectammina perplexa was described from the Middle Eocene Lodo Formation of California by
Israelsky (1951), and later Martin (1964) used this term for his specimens form the Paleocene Moreno
Formation. The test was described as being "smooth, minutely granular, with largely siliceous cement".
Israelsky placed the specimens described as S. clotho by Cushman (1934), Cushman & Jarvis (1928)
and Cushman & Renz (1946) in the synonymy of his S. perplexa. The type specimens preserved in the
Cushman Collection (USNM 560501 & USNM 560502: see Fig. 104-4) are strongly compressed and
decrease in width distally. The mean thickness of the two preserved specimens is only 0.07 mm. It is the
thinnest end member of the named synonyms of S. spectabilis measured by Kaminski (1984). Such
relatively thin forms have been given the name "S. spectabilis forma perplexa" by Kaminski (1984).
Workers familiar with the species S. spectabilis from the North Sea region have noticed that the Eocene
morphotype is apparently smaller and thinner than the Paleocene morphotype (e.g., King, 1989). Gradstein
(1983) regarded the transition from robust Paleocene forms to thinner Eocene forms to reflect an
evolutionary transition. We must point out, however, that flat, thin forms that are not significantly different
in overall dimensions from the Eocene type specimens of S. perplexa are also found in the mid-Paleocene
Moreno Shale of California. For this reason, Kaminski (1984) regarded this flattened morphotype as simply
an ecophenotype of S. spectabilis, and assigned it an informal name (S. spectactabilis forma perplexa),
following the practise of Feyling-Hansen (1972) for distinct ecophenotypic variants. However, Charnock &
Jones (1990) regarded this form as being "no more than infrasubspecifically distinct from the typical form".
It is possible that at least in the Paleocene, the paleobathymetric setting influences the shape of S.
spectabilis, with the thicker, more robust forms occurring more frequently in deeper assemblages. Eocene
topotype specimens from the Polish Carpathians, which are from undoubtedly deep-water assemblages, are
significantly thicker (mean = 0.18 ± 0.3 mm) than the types of S. perplexa.
Spiroplectammina mexiaensis was originally described by Lalicker (1935) from the Mexia clay member
of the Wills Point Formation in Mexia Co., Texas. Three slides of specimens are preserved in the Cushman
Collection. The holotype (CC21924) is a megalosphaeric specimen, and because the test is white, not
silicified, the exact arrangement of the early chambers is not visible. The biserial part consists of nine pairs
of chambers with depressed sutures. The holotype is 0.6 mm in length and 0.22 mm in maximum width.
The two paratypes housed in the collection (CC21925, USNM 371544) are also megalosphaeric forms. All
of the megalosphaeric syntypes possess spiral parts that are narrower than the widest portion of the biserial
part.
Two species of Spiroplectammina originally described from South America considered here to be
junior synonyms of are S. grzybowskii Frizzell, and S. brunswickensis Todd & Kniker (Fig. 104-4). Frizzell
(1943) originally described S. grzybowskii from ?Late Cretaceous of a well drilled near Negritos Peru (J.P.
well 1755, 2310'). This fauna is in fact Danian in age (see remarks under R. garcilassoi). In his description
of the species, Frizzell placed in synonymy the specimens from Lizard Springs illustrated by Cushman &
Jarvis (1928) as S. clotho. Cushman (1947) later reported S. grzybowskii from additional localities in
 434

Trinidad and Venezuela, but remarked that its relationship to his S. clotho is "unclear". The paratypes of S.
grzybowskii preserved in the Cushman Collection (CC39551 - CC39552) are relatively large in comparison
to forms described from North America, but its dimensions are not significantly different from those of the
topotype specimens of S. spectabilis from the Polish Carpathians.
The second South American species is S. brunswickensis, described by Todd & Kniker (1952) from the
Upper Eocene Aqua Fresca Shale of southern Chile. Todd & Kniker stated that the species "differs from S.
directa in its more elongate test and lack of a keel" but added that the two species "seem to be closely
related and may prove indistinguishable". The holotype of S. brunswickensis (CC64279) is a
megalosphaeric specimen 0.9 mm in length, 0.22 mm in width, that has 13 pairs of chambers in its biserial
part. Paratype soecimens (CC64280) possess spiral portions that are generally narrower in diameter than the
maximum width of the biserial part.

Text-fig. 104-4. Type specimens of four species from the Cushman Collection considered here to be junior synonyms of S.
spectabilis (all camera lucida drawings by MAK, scale bar = 0.5 mm). 1-2. Spiroplectammina perplexa Israelsky, Lodo Formation,
Tumey Hills, Fresno Co. CA. 1. Holotype (USNM 560501); 2. Paratype (USNM 560502). 3-4. Spiroplectammina brunswickensis
Todd & Kniicker, middle Agua Fresca Formation, Brunswick Peninsula, Magallanes Province, Chile. 3. Holotype (CC 64279); 4.
Paratype (CC 64280); 5-6. Spiroplectammina grzybowskii Frizzell, Mel Paso Shale, I.P. Co well 1755, 1960', paratypes (CC 39551,
CC 33552); 7-9. Spiroplectammina mexiaensis Lalicker, Mexia Point Formation, "2 mi N & SW of the center of Mexia, Limestone
Co., TX", 7. Holotype (CC 21924); 8-9. Paratypes (CC 21925; USNM 371544).

The generic affiliation of S. spectabilis is still unsettled due to the uncertain status of the genus
Bolivinopsis. According to the current generic definitions of Loeblich & Tappan (1987), S. spectabilis
clearly belongs in the genus Bolivinopsis. The type species of Spiroplectammina, (Textularia agglutinans
var. biformis Parker & Jones, 1865), has organic cement, a rounded periphery, and an initial spiral part that
is of lesser width than the biserial portion of the test. We have previously regarded the genus Bolivinopsis
as nomen dubium following the suggestion of Frizzell (1943) because of the lack of preserved specimens of
its type species, Bolivinopsis capitata Yakovlev, 1891. This species was reported from the Upper
Cretaceous of Russia by Yakovlev, and the nature of its cement is unknown. Macfadyen (1933) considered
B. capitata to be a junior synonym of Spiroplecta rosula Eherenberg, a species with an imperforate test and
calcareous cement. Kisselman (1964) has pointed out that even in the type region of B. capitata, the
designation "B. rosula" is used for these forms. Loeblich & Tappan (1987) remarked that "Cenozoic species
previously assigned to Bolivinopsis are probably not congeneric", and listed the genus as restricted to the
Late Cretaceous. They do not mention the composition of the cement, and Bolivinopsis is distinguished
from Spiroplectammina in being more elongate and compressed, and in having a smoothly finished wall.
Although we agree in principle with the generic scheme for the spiroplectamminids adopted by Loeblich &
Tappan, in our opinion, the genus Bolivinopsis needs to be validated by examining the nature of its cement
and designating a lectotype for its type species. Until this is done, we retain the Cenozoic forms in
Spiroplectammina.

ILLUSTRATIONS: Plate 104 - Spiroplectammina spectabilis (Grzybowski) emend. Kaminski –

 435
Plate 104 - Spiroplectammina spectabilis (Grzybowski) emend. Kaminski

Type specimens of S. spectabilis and Grzybowski's junior synonyms. Fig. 1a-c. "Spiroplecta spectabilis
Grzybowski" (Grzybowski Collection; UJ-132-P 1/82a) Potok H-33 well, 170 m, probably the plesiotype
of Grzybowski (1898, pl. 12, fig. 12). 1a,b. reflected light, 1c. cross-polarised light, Lectotype, x56; Fig.
2a,b. "Spiroplecta spectabilis Grzybowski" (Grzybowski Collection; UJ-132-P 1/82b), "Potok" exact
locality unknown, 2a,b. reflected light, x50; Fig. 3a-c. "Spiroplecta brevis Grzybowski" (Grzybowski
Collection; UJ-132-P 1/83a) "Potok" exact locality unknown, plesiotype of Grzybowski (1898, pl. 12, fig.
13), 3a,b. reflected light, 3c. cross-polarised light, x66; Fig. 4a,b. "Spiroplecta foliacea Grzybowski"
(Grzybowski Collection; UJ-132-P 1/84a) Potok H-40 well, 24 m, 4a. reflected light, 4b. cross-polarised
light, x74; Fig. 5a-c. "Spiroplecta clotho Grzybowski" (UJ-133-P 2/100a) Bartne 14 (80), plesiotype of
Grzybowski (1901, pl. 7, fig. 18), 5a,b. reflected light, 5c. cross-polarised light, x69; Fig. 6a,b.
"Spiroplecta costidorsata Grzybowski" (UJ-132-P 1/85b) Potok H-26 well, 96 m, probably the plesiotype
of Grzybowski (1898, pl. 12, fig. 11), 6a. reflected light, 6b. cross-polarised light, x78. All specimens
from the Grzybowski Collection, courtesy of S. Geroch (light microscope photos).
 436

ATLAS SP. # 105

Spiroplectammina trinitatensis Cushman & Renz, 1948

Fig. 105-1. Type figures of Spiroplectammina trinitatensis,

from Cushman & Renz (1948)

ORIGINAL DESIGNATION: Spiroplectammina trinitatensis Cushman & Renz, 1948.

TYPE REFERENCE: Cushman, J.A. & Renz, H.H. 1948. Eocene foraminifera of the Navet and Hospital Hill formations of
Trinidad, B.W.I. Contributions from the Cushman Laboratory for Foraminiferal Research Special Publication, 24, p. 11, pl. 2, figs.
13-14.

TYPE SPECIMEN: Housed in the microfossil collections of the US Natural History Museum, Smithsonian
Institution, Washington, D.C. Holotype is registered as CC 57395.

TYPE LEVEL: Late Eocene, Hospital Hill Formation, San Fernando Group.

TYPE LOCALITY: Northwest slope of Hospital Hill, at the San Fernando Hospital, east side of the road
from San Fernando to Point Bontour, Trinidad.

DIAGNOSTIC FEATURES: Differs from Spiroplectammina spectabilis in its greater dimensions and
disproportionally greater thickness along the medial line. In addition, the later chambers are higher than in
S. spectabilis, and sutures in the biserial part are more strongly incised. The peripheral margin is more
irregularly dentate.

SIZE: Cushman & Renz reported specimens 2.0 mm in length, 0.5 mm in width.

SYNONYMS: None verified.

OBSERVED OCCURRENCES: This species was originally described from the Navet Formation of
Trinidad by Cushman & Renz (1948). The age of Cushman's sample from the type locality was determined
to be earliest Late Eocene (Zone P15/NP18) by McCabe et al. (1993). Bermúdez & Gamez (1966) reported
it from the Middle Eocene Punta Mosquita and El Dátil Formations of Isla Margarita, Venezuela, in
assemblages dominated by calcareous benthic foraminifera with the deep-water index species Nuttallides
truempyi. Beckmann (1994) listed it from Late Paleocene (Zone P5) in the Lizard Springs Formation of
Trinidad. Müller-Merz & Oberhansli (1991) illustrated a specimen from Late Eocene (Zone P17) of DSDP
Hole 522 in the South Atlantic. Bolli (1994) reported its stratigraphic range in Trinidad as Middle to Late
Eocene (Zone P10 to P16). We have observed it in the Middle Eocene Navet Formation of Trinidad and in
the Middle to Late Eocene of ODP Hole 647A, Labrador Sea.

KNOWN STRATIGRAPHIC RANGE: Middle to Late Eocene (P10 - P16).

 437

BATHYMETRY: Bathyal to abyssal

REMARKS: In an early study of the type specimens of Spiroplectammina trinitatensis, Kaminski (1984)
concluded that this species is of significantly larger dimensions than any other Cenozoic species of
Spiroplectammina housed in the Cushman Collection. The use of the trinomial nomenclature was
suggested, and large Middle Eocene forms were assigned to S. spectabilis forma trinitatensis.

-200 -200
S. spectabilis
S. spectabilis S. trinitatensis
S. trinitatensis
-300 -300

-400 -400

-500 -500

-600 -600

-700 -700
0.0 0.1 0.2 0.3 0.4 0.2 0.4 0.6 0.8 1.0 1.2 1.4
Width of initial spire (mm) Length of test (mm.)

Fig. 105-2. Measurements of test length and diameter of the initial spire of megalosphaeric forms of S. spectabilis and S.
trinitatensis in samples from the Middle-Upper Eocene at ODP Hole 647A.

However, at ODP Site 647, this robust form occurs alongside forms that are significantly smaller. Our
measurements of all the Spiroplectammina specimens from this site reveals that there are two distinct
populations (Fig. 105-2). The presence of both large and small forms co-occuring at this locality indicates
genetic isolation, and we therefore regard S. trinitatensis as a species distinct from S. spectabilis.
Spiroplectammina bukkiana Sztrákos, 1979, described from the upper part of the Kiscell Clay (Early
Oligocene) in Hungary appears to be a related species. Its affiliation to S. trinitatensis needs to be
established by further study.

ILLUSTRATIONS: Plate 105 - Spiroplectammina trinitatensis Cushman & Renz –

 438
Plate 105 - Spiroplectammina trinitatensis Cushman & Renz

Figs. 1a–2b. Upper Eocene (P16), San Fernando Group, Trinidad. "Hospital Hill" locality of Cushman &
Renz (1946), Topotypes from Cushman's sample residues, 1a,b, x50; 2a,b, x65; Fig. 3a,b. Middle Eocene
(P13), Navet Formation Trinidad. "Penitence Hill marl" locality of Cushman & Renz (1946), Topotype
from Cushman's sample residues, x52; Fig. 4a-c. Middle Eocene, ODP Site 647, southern Labrador Sea.
Sample 647A-43R-3, 104-108 cm, Microsphaeric specimen, 4a,b. x87, 4c. detail of aperture, x500; Figs.
5–6. Middle Eocene, ODP Site 647, southern Labrador Sea, sample 647A-43R-3, 104-108 cm.
Megalosphaeric specimens, 5. x105, 6. x115; Fig. 7a-c. Middle Eocene, ODP Site 647, southern Labrador
Sea, sample 647A-43R-1, 95-98 cm. Microsphaeric specimen; 7a,b. x90, 4c.detail of aperture, x310.
 439

ATLAS SP. # 106

Spiroplectinella dentata (Alth, 1850)

Fig. 106. 1. Type figure of "Textularia dentata" from Alth

(1850); 2. Topotype specimen from Lvov illustrated by
Cushman (1932), here designated the neotype.

ORIGINAL DESIGNATION: Textularia dentata Alth, 1850.

TYPE REFERENCE: Alth, A. 1850. Geognostische-paläntologische Beschreibung der nächsten Umgebung von Lemberg.
Abhandlungen Naturwissenschaften Wien, 3, p. 262, pl. 13, fig. 13. see also: Cushman, J.A. 1932. Textularia and related forms
from the Cretaceous. Contributions from the Cushman Laboratory for Foraminiferal Research, 8, p. 91, pl. 11, fig. 7a,b [neotype].

TYPE SPECIMEN: Not originally designated. The Alth collection is presumed lost. A specimen from the
Lvov Marls illustrated by Cushman (1932) is preserved in the Cushman Collection (CC 17595). We
designate this specimen the neotype.

TYPE LEVEL: Maastrichtian, Lvov Marls.

TYPE LOCALITY: Outcrops of marls near the city of Lvov (Lemberg) in the western Ukraine.

DIAGNOSTIC FEATURES: Test initially planispiral, later biserial, rhomboidal in cross-section. Initial
portion increases rapidly in width, later more gradually or not at all. Periphery is often dentate. Chambers
are relatively high, with approximately 11 pairs in the biserial part. Sutures in the initial portion of the
biserial part are straight, and become more arched towards the distal end. As a result, the apex of larger
specimens is more rounded in outline. Megalosphaeric and microsphaeric forms differ markedly in the
dimensions of the initial spire and in the total number of chambers. Megalospheric forms are shorter, with a
more broadly rounded initial portion, and more parallel sides. Wall imperforate, agglutinated with
calcareous cement. Aperture interiomarginal, a low slit.

SIZE: Topotype specimens from Lvov are up to 0.9 mm in length; specimens from the Carpathian flysch
attain a length of 1.4 mm.

SYNONYMS: None verified.

OBSERVED OCCURRENCES: Although S. dentata was first described from marly sediments deposited in
a neritic setting (the Maastrichtian Lvov marls), this designation has also been applied to deep-water forms
from flysch-type assemblages (e.g., Cushman & Jarvis, 1932; Grün, 1969). It is widely distributed in slope
assemblages from marly sediments of the continental margins, in Scaglia-type assemblages from deep-
water limestones, and in lesser numbers from flysch-type assemblages from turbidite units (such as Lizard
Springs), though it may be redeposited. In its type area (the Volynsko-Podolska Platform in the
 440
Plate 106 - Spiroplectinella dentata (Alth)

Figs. 1a–3b. Maastrichtian, topotype specimens from the Lvov Marls; 1a,b. largest microspheric form in
our collection with slightly arched final chambers, x70; 2a,b. smaller microspheric form, x95; 3a,b.
megalospheric form, x95; Fig. 4a,b. Maastrichtian, Dydnia Poland, Skole Unit of the Carpathians,
microspheric form, x67; Figs. 5–6. Danian, Gossau beds of the Bavarian Alps, Wasserfallgraben section,
5. microspheric form, x60, 6. megalospheric form, x90; Fig. 7. Danian, Lizard Springs Formation of
Trinidad, x95; Fig. 8. Campanian, Viking Graben of the North Sea, Total 3/25-1 well, 2840 m, x95.
 441

Ukraine), Kaptarenko-Chernousova et al. (1979) reported its range as late Campanian to Maastrichtian.
Kavary & Frizzell (1963) reported it as Spiroplectammina dentata from the Lower Paleocene of the Zagros
Mountains in Iran, and Braga et al. (1975) recorded it from the Paleocene (P1c–P5) of the Possagno section
in Italy.
We have observed it in the Danian of the Lizard Springs Formation of Trinidad, in the Danian of
Zumaya Spain, in the Campanian to Danian of the Gubbio Section of Central Italy, and in the Paleocene of
the Gosau Beds in the Austrian and Baverian Alps. It is widely distributed in the Campanian to
Maastrichtian of the Carpathian flysch. On the Labrador Margin it occurs in low numbers in the
Maastrichtian to Paleocene. In the Central North Sea and offshore mid Norway it is found with rare
specimens in Middle to Late Campanian (Tritatia dubia Zone, Gradstein et al., 1999).

KNOWN STRATIGRAPHIC RANGE: Campanian to Paleocene.

BATHYMETRY: Outer neritic to lower bathyal.

REMARKS: The type figure of Textularia dentata illustrated by Alth (1850) depicts a megalospheric
individual. The specimen illustrated by Cushman (1932) in his pl. 11, fig. 7a,b is a microspheric form. Both
forms occur in our topotype material from the Lvov marls. Topotypes from the Lvov marls are white in
colour, are thickest at the terminal end, and have a relatively flat apertural face. However, specimens from
bathyal deposits attain a larger size and have more arched final chambers than the topotypes (e.g.,
specimens illustrated in Plate 106, figs. 4-5). These deep-water forms have a more tapered outline (the
thickest part of the test is not necessarily across the final chambers) and are usually more strongly dentate
than the topotypes from Lvov. Specimens such as these have been illustrated as S. dentata by Cushman &
Jarvis (1932) and by Grün (1969). Carpathian workers informally refer to this morphotype as "S. dentata
sensu Grün, 1969". For this reason, we initially regarded this species as a plexus, and used the designation
"S. ex gr. dentata" to describe the deep- water forms. We have not observed the deep-water morphotype
from strata younger than Paleocene age. Specimens in the Cushman Collection from the Eocene Navet
Formation are triangular in outline and belong in a different species. We place this species in
Spiroplectinella in accordance with the generic classification of Loeblich & Tappan (1987).

ILLUSTRATIONS: Plate 106 - Spiroplectinella dentata (Alth) –

 442

ATLAS SP. # 107

Spiroplectinella israelskyi (Hillebrandt, 1962)

Fig. 107. 1. "Spiroplectammina sp. A" from Israelsky (1951); 2-3. Spiroplectammina israelskyi from Hillebrandt (1962).

ORIGINAL DESIGNATION: Spiroplectammina israelskyi Hillebrandt, 1962.

TYPE REFERENCE: Hillebrandt, A. 1962. Das Paleozän und seine Foraminiferen-fauna im Becken von Reichenhall und
Saltzburg. Abhandlungen, Bayerische Akademie der Wissenschaften, Mathematisch-Naturwissenschaftliche Klasse, 108, p. 30, pl.
1, figs. 5a-7b.

TYPE SPECIMEN: Deposited in the Protozoan Collection of the Bavarian State Museum in Munich. The
holotype is registered as PROT #1136.

TYPE LEVEL: Paleocene, Gosau Unit of the Alps.

TYPE LOCALITY: Eitelgraben creek in the Untersberg Nature Park, on the northern flank of the
Untersberg, near Wegscheid Austria. The locality has been described in greater detail in the IWAF-3
excursion guidebook (Kuhnt et al., 1989).

DIAGNOSTIC FEATURES: Test initially planispiral, later biserial, lanceolate in outline, rhomboidal in
cross-section. The spiral portion consists of 4-5 chambers, with the biserial part having as many as 13 pairs
of chambers. The initial portion initially increases rapidly in width, but in the later part of the biserial
portion the chambers can vary in width. As a result, the outline is slightly irregular. Megalospheric forms
have a distinct spiral portion, >0.1 mm in diameter, with four chambers. In microspheric forms the initial
spire is usually indistinct. The wall is imperforate, built of both calcareous and noncalcareous grains, with
much cement, and has a smooth finish. Aperture interiomarginal.

SIZE: Type specimens attain a length of 1.35 mm.

SYNONYMS:
Spiroplectammina sp. A. Israelsky, M.C., 1951. US Geol. Surv. Prof. Paper, 240-A, p. 13, pl. 3, figs. 17-19
[Paleocene, California; by original designation].

OBSERVED OCCURRENCES: As Spiroplectammina sp. A, Israelsky (1951) reported this species from
the Paleocene of the Lodo Formation of California. Hillebrandt (1962) described it as S. israelskyi from the
mid-Paleocene of the Eitelgraben section of the Austrian Alps. These are marly sediments of the Alpine
Gosau Unit that were deposited at upper bathyal paleodepths. De Klasz & De Klasz (1990) illustrated a
specimen as S. dentata from Danian marls of the Ultrahelvetic Zone in Bavaria. This species occurs
sometimes abundantly in the Campanian-Paleocene of the Scaglia-type assemblages in the Gubbio Sections
of Italy (Kuhnt, 1990). Widmark (1997) reported it from the Maastrichtian at Sites 525 and 527 in the South
Atlantic. Peryt et al. (1997) observed the FO of S. israelskyi in Zone P1a (Danian) in flysch sediments of
the Rotwandgraben section, Eastern Alps (Austria). Milner (1997) illustrated specimens as
 443
Plate 107 - Spiroplectinella israelskyi (Hillebrandt)

Fig. 1. Mid Paleocene (Zone P3) Eitelgraben, Gosau Zone, Austria, Topotype, x95; Fig. 2. Lowermost
Paleocene, “K/T boundary clay”, Monte Conero, Italy, x82; Fig. 3. Lower Paleocene, Zumaya Spain, x52;
Figs. 4-7. Lowermost Paleocene, “K/T boundary clay”, Sopelana Spain, 4. x51, 5. x55, 6. x70, 7. x54;
Figs. 8-11. Maastrichtian, Gubbio Italy, 8. x96, 8, x102, 10. x100, 11. x132.
 444

S. dentata from the Paleocene Moogli Mudstone and Burns Peak Beds of Papua New Guinea. We have also
observed it in the Campanian of the Subsilesian Unit of the Polish Carpathians, in the Paleocene of the
Zumaya section of northern Spain, in the Maastrichtian of northern Morocco, and in the Campanian-
Maastrichtian in ODP Holes 398D and 543A in the North Atlantic.

KNOWN STRATIGRAPHIC RANGE: Campanian to Paleocene.

BATHYMETRY: Bathyal.

REMARKS: This species can be distinguished from S. dentata by its smooth finish, small initial spire, and
irregularly parallel sides. Specimens typically display irregular or nonuniform chamber growth, and as a
result the peripheral margin may have invaginations. In addition, S. israelskyi normally has an orange
colour, which is probably owing to its abundant calcareous cement.

ILLUSTRATIONS: Plate 107 - Spiroplectinella israelskyi (Hillebrandt) –

ATLAS SP. #108

Spiroplectinella subhaeringensis (Grzybowski, 1896)

Fig. 108-1. Type figures of "Textularia subhaeringensis", from Grzybowski (1896).

ORIGINAL DESIGNATION: Textularia subhaeringensis Grzybowski, 1896 var. α; and Textularia

subhaeringensis Grzybowski, 1896 var. β.

TYPE REFERENCE: Grzybowski, J. 1896. Otwornice czerwonych iłów z Wadowic. Rozprawy Wydziału Matematyczno-
Przyrodniczego, Akademia Umiejętności w Krakowie, serya 2, vol. 30, p. 285, pl. 9, figs. 13, 16. See also: Liszka, S. & Liszkowa, J.
1981. Revision of J. Grzybowski's paper (1896) "Foraminifera of the red clays from Wadowice". Rocznik Polskiego Towarzystwa
Geologicznego, 51, 153-208. pl. 3, figs. 6a-7b.

TYPE SPECIMEN: Not originally designated. The specimens coresponding to Grzybowski's illustrations
were found by Liszka & Liszkowa (1981), who regarded the megalosphaeric specimen described as "T.
subhaeringensis Grzybowski, 1896 var. α " as the holotype. These specimens are housed in the
micropaleontological collections of the Jagiellonian University, Kraków (Grzybowski Collection slide nos.
UJ-115P, 44, 47)

TYPE LEVEL: Campanian, Subsilesian Unit of the Polish Carpathians.

TYPE LOCALITY: See Hyperammina dilatata.

DIAGNOSTIC FEATURES: Test initially planispiral, later biserial, nearly as broad as long, rhomboidal in
cross-section. The spiral portion consists of 4-5 chambers, with the biserial part having as many as 17
chambers. The initial portion increasing rapidly in width and thickness. The initial part of the test is
 445

triangular in outline, with the sides diverging at an angle of 70-105°; the last two chambers are broadly
rounded in outline. The periphery in the initial part is acute and serrate. Sutures in the biserial part are
arched and raised slightly. Wall imperforate, with calcareous cement. Aperture an interiomarginal slit.

SIZE: Topotype specimens range up to 0.9 mm in length (average 0.6 mm).

SYNONYMS:
Textularia excolata Cushman. Cushman, J.A., 1926. Bull. Am. Assoc. Petr. Geol. 10, 585; pl. 15, fig. 9
[Paleocene, Mexico].

OBSERVED OCCURRENCES: Grzybowski (1896) originally described T. subhaeringensis from

Campanian red clays from Wadowice, Poland. As Spiroplectammina subhaeringensis, the species has also
been reported from other Upper Cretaceous and Paleocene beds from the Carpathians in Poland (Huss,
1966; Szczechura & Pożaryska, 1974; Morgiel & Olszewska, 1981; Geroch & Nowak, 1984, Bąk 2004),
and Romania (Neagu, 1970). Tjalsma & Lohmann (1982) recorded it from the Paleocene DSDP Site 21 in
the South Atlantic. As Spiroplectammina excolata, it was described from the Paleocene Velasco Formation
of Mexico (Cushman, 1926) and Lizard Springs Formation of Trinidad (Cushman & Jarvis, 1932).
Hillebrandt (1962) reported it from the Paleocene Gossau Beds of the Austrian Alps, and Beckmann et al.
(1982) recorded it from Middle Maastrichtian in the Pianno di Brenno Formation at Monte Giglio, Italy.
Beckmann (1994) recorded it as Spiroplectinella excolata from Zones P1d to P5, Paleocene of Trinidad.
We have observed this species throughout the Polish Carpathians, in the Campanian of DSDP Hole
543A in the western North Atlantic, in the Paleocene of the Lizard Springs Formation of Trinidad, in the
Upper Cretaceous Gibraltar flysch of southern Spain, and in the Paleocene Kamalapuram Formation of the
Cauvery Basin, India.

KNOWN STRATIGRAPHIC RANGE: Campanian to Paleocene.

BATHYMETRY: Bathyal to abyssal.

REMARKS: This species is readily distinguished from other species of Spiroplectinella by its rapidly
increasing width and thickness and by its raised arched sutures. Paleocene forms described as T. excolata
sometimes develop slightly irregular sutures in the adult portion of the test. We agree with the suggestion of
Szczechura & Pożaryska (1974) in regarding this species as a junior synonym of S. subhaeringensis.
Grzybowski (1896) originally described two varieties of T. subhaeringensis. He described the smaller
alpha variety as follows:

"Test composed of low, arcuate chambers, pointing backward toward the initial end of the test. The small early chambers
widen rapidly, alternating obliquely and overlapping at the sides, so that the test forms, in outline, a semicircle at the
terminal end and a blunt triangle at the inital end. Ridges run along the sutures toward the sharp periphery, where they end
in prominent spines slanting toward the initial end, so that the periphery is serrate. Aperture at the base of the last
chamber, an oval slit."

Grzybowski separated a second variety of T. subhaeringensis, which he called the "beta" variety, based on
differences in the outline. He described it as :

"Test similar in structure to the preceding form, but more compressed and elongate, lanceolate in outline, initially more
rounded. Sutural ridges form less elongate spines at the periphery."

Grzybowski's description of the beta variety gives the impression that the initial spiral portion is larger.
Liszka & Liszkowa (1981) considered the shorter and wider alpha variety to represent the megalospheric
generation, whereas they regarded the beta variety to be the microspheric form.
We measured 48 topotype specimens of S. subhaeringensis in a effort to determine whether it is
possible to discriminate between the two described varieties, and whether they can be attributed to different
 446

generations. The following parameters were measured in transmitted light: length; thickness; maximum
width of the biserial part; diameter of the proloculus; and the width of the initial spire across the proloculus
(perpendicular to the long axis of the test). We measured all specimens, without preselecting "alpha" and
"beta" forms. The data are presented in Figures 108-2a,b.

0.8
0.75 Max width
Max thickness
0.65 0.6
0.55
0.4
0.45

0.35
0.2
0.25 Max width
a Width Spire b
0.15 0.0
0.2 0.4 0.6 0.8 1.0 0.04 0.05 0.06 0.07 0.08 0.09 0.10
Length (mm) Proloculus diameter (mm)
Figure 108-2. (a) measured growth-related parameters of topotype specimens of S. subhaeringensis. (b) relationship between
proloculus diameter, width of the initial spire, and maximim width of the biserial part. This is a measure of the "lanceolate" versus
"flairing" outline of the test.

As specimens increase in length, the rate of increase in both width and thickness decreases
logarithmically (Figure 108-2a). Among larger individuals, the rate of increase in thickness is slower than
the rate of increase in width, so larger specimens appear to be more flattened in their later chambers.
Typical "alpha" forms, in the sense of Grzybowski, are thicker, more flairing ones. Among our topotype
specimens, the test becomes more "lanceolate" after the 13th chamber, as the rate of increase in width and
thickness decreases.
If the differences in outline described above can be attributed to alternate generations, then one
generation ought to display a greater difference between the maximum width of the biserial part and the
width of the test across the spire. Figure 108-2b shows the regression lines between proloculus diameter
and the two width parameters, and figure 108-3 presents the distribution of the proloculus diameter
measurements.

12

10

0
0.04 0.05 0.06 0.07 0.08 0.09 0.10
Proloculus diameter (mm)

Figure 108-3. Distribution of measurements of proloculus diameter among 48 topotypes of S. subhaeringensis.

 447
Plate 108 - Spiroplectinella subhaeringensis (Grzybowski)

Fig. 1a-c. Lectotype from the Grzybowski Collection; Fig. 2a-c. Syntype from the Grzybowski
Collection; Fig. 3a,b. Paleocene, Gulf of Mexico, Eurica Core E68-151A, x215; Fig. 4a,b. Paleocene,
Lizard Springs Formation of Trinidad, x80; Fig. 5a,b. Paleocene, Eitelgraben, Austria, x120.
 448

There appears to be no clear relationship between proloculus diameter and the width of the spire, but there
is a very weak inverse relationship between proloculus diameter and the width of the biserial part. The
largest difference between the two width parameters is observed among individuals with the smallest
proloculus diameter. There apparently is a weak relationship between test shape and proloculus diameter,
but these differences are not significant. There is also no relationship between proloculus diameter and the
total number of chambers. Moreover, the distribition of our measurements of the proloculus diameter is
unimodal (Fig. 108-3). Therefore, we cannot confidently distinguish between megalospheric and
microspheric generations even though the diameter of the proloculus varies by a factor of two. We conclude
from this analysis that Grzybowski's two varieties may reflect differences in ontogeny, with the juvenile
forms representing the more flairing "alpha" variety. This analysis does not uphold Grzybowski's
subdivision of S. subhaeringensis into two varieties.
Grzybowski noted that T. subhaeringensis is similar to the initial portion of the species Bigenerina
capreolus (d'Orbigny) illustrated by Brady (1884) in pl. 45, fig. 3 (=Vulvulina pennatula according to
Barker, 1960). He remarked however,

"because among several dozen specimens [...] I did not find a single one which showed the least trace of uniserial chamber
arrangement, I cannot indentify this form with Brady's [...]. It was necessary, therefore, to separate this form as a distinct
species.".

Grzybowski believed that T. subhaeringensis may be ancestral to the Recent species Vulvulina pennatula, a
claim that we do not dispute.

ILLUSTRATIONS: Plate 108 - Spiroplectinella subhaeringensis (Grzybowski) –