Type species: Globigerina (Eoglobigerina) eobulloides Morozova, 1959, emended.

Diagnostic characters: Low, trochospiral test with 10-16 chambers, 4-6 1/2 globular chambers in the ultimate whorl. The trochospire is moderately to highly elevated; aperture is interiomarginal, umbilical to slightly extra-umbilical, a low, rounded arch bordered by a thin, narrow lip; and umbilicus is small and is open to the apertures of the surrounding chambers. The wall is cancellate and spinose with the spine holes situated along the cancellate ridges.

Discussion: Hemleben et al. (1991) demonstrated that Eoglobigerina had a spinose morphology that separates it from other cancellate forms in the Danian that are nonspinose. The concept of Eoglobigerina followed here is similar to that of previous workers except that it is emended to include the spinose character.

Globigerina edita Subbotina, 1953, p. 62, pl. 2, figs. 1a-c [(Zone of rotaliform Globorotalia (Danian Stage), Kuban River section, North Caucasus)].-Fox and Olsson, 1969, p. 1398, pl. 168, figs. 1-4 (lower Paleocene, Cannonball Fm., North Dakota).-Shutskaya, 1970b, pl. 18, figs. 14a-c (non 12a-c) (middle Subzone of Gl. trivialis-Gl. conusa-Gl. compressa Zone, Malyi Balkhn Ridge, W. Turkmenia).

Globorotalia (Globorotalia) edita (Subbotina). -Hillebrandt, 1962, p. 130, pl. 11, figs. 14, 15 (Zone A, lower Paleocene, Reichenhall-Salzburg Basin, Austro-German border).

Eoglobigerina edita edita (Subbotina). -Blow, 1979, p. 1210, pl. 61, figs 2, 3; pl. 66, fig. 1 (Zone P , DSDP Hole 47.2/11/3: 148-150 cm, Shatsky Rise, NW Pacific Ocean); pl. 69, fig. 6 (Zone P1, DSDP Hole 47.2/11/3: 0-5 cm); pl. 72, figs. 6, 8 (Zone P1, DSDP Hole 47.2/11/3: fig. 6, 148-150 cm; fig. 8, top section); pl. 79, fig. 3 (Zone P2, DSDP Hole 20C/6/4: 72-74 cm, South Atlantic Ocean).

Eoglobigerina edita (Subbotina). -Hemleben, M¸hlen, Olsson and Berggren, 1991, p. 126, pl. 7, fig. 4-6 (Zone P , Clayton Basal Sands, Millers Ferry, Alabama). -Berggren, 1992, p. 562, pl. 1, figs. 5, 6 (Zone P1b, ODP Hole 747A/19H/CC, Kerguelen Plateau, Southern Indian Ocean). -Olsson, Hemleben, Berggren and Liu, 1992, p. 197, pl. 2, figs. 1-5, 7 (Zone P , fig. 1-4, Clayton Basal Sands; Zone P1a, figs. 5, 7, Pine Barren Mbr., Clayton Fm., Alabama).

Globigerina edita Subbotina VAR. polycamera Khalilov, 1956, p. 235, pl. 1, figs. 1a-c (Danian Stage, Akhchakuima, NE Azerbaizhan).-Shutskaya, 1970a, p. 119, pl. 18, figs. 11a-c (middle Subzone of Gl. trivialis-Gl. daubjergensis-Gl. compressa Zone, Khazin-Don River section, N Caucasus).

Eoglobigerina edita polycamera Khalilov. -Olsson, Hemleben, Berggren and Liu, 1992, p. 197, pl. 2, fig. 6 (Zone P1a, Pine Barren Member, Clayton Fm., Millers Ferry, Alabama).

Globigerina (Eoglobigerina) hemisphaerica Morozova, 1961, p. 11, pl. 1, fig. 4 (Zone Dn1 II, Tarkhankut Peninsula, Crimea; lower substage of Danian Stage).

Globigerina (Eoglobigerina) tetragona Morozova, 1961, p. 13, pl. 1, fig. 2 (Zone Dn1 I, Precaspian Basin, Novouzensk, lower substage of Danian Stage).

Globigerina (Eoglobigerina) pentagona Morozova, 1961, p. 13, pl. 1, fig. 3 (Zone Dn1 I, Tarkhankut Peninsula, Crimea; lower substage of Danian Stage).

Globigerina (Eoglobigerina) theodosica Morozova, 1961, p. 11, pl. 1, fig. 6 (Zone Dn1 I, Tarkhankut Peninsula, Crimea; lower substage of Danian Stage).

Original description: Shell small, rounded, with tall spiral consisting of 2 1/2-3 whorls. The last whorl is made up of
4 1/2-5 spherical chambers, usually of almost equal size. Peripheral margin rounded, scalloped. The most characteristic feature of this species is the turret-like dorsal surface which is associated with the large size of the chambers in the earlier whorls.

The ventral side is feebly convex compared with the dorsal. The umbilicus is small and shallow but distinct. The chambers of the early whorls are compactly arranged and closely packed together, those of the last whorl are arranged much more freely giving a scalloped appearance to the peripheral margin. Sutures deep, slightly curved. The orifice has the form of a small slit which extends along the marginal suture. Walls smooth, with small pores. Mean dimensions: diameter 0.25 mm, thickness 0.15 mm.

Diagnostic characters: The species is characterized by a moderately high trochospiral test with 4 1/2 to 5 (occasionally up to 7) globular chambers which increase gradually in size in the ultimate whorl, and a strongly lobulate peripheral margin. The aperture is a rounded umbilical to slightly extraumbilical aperture which is bordered by a narrow lip. The umbilicus is small but open to the previous chambers. The cancellate wall is weakly developed and spinose with numerous spine holes along the cancellate ridges. The overall size of the test is generally less than 250 um.

Discussion: Considerable variation in height of the initial coil and the position of the aperture from umbilical towards an extraumbilical position occurs in this species. Blow (1979) discussed the high degree of variability among early eoglobigerinids and the apparent fact that few morphotypes exhibit a consistent expression of morphologic characters. Thus, he cautioned that strict morphologic limits cannot be as rigidly applied to early eoglobigerinids as to the later, more stable morphospecies of globigerinacean genera. Blow also distinguished as a subspecies of edita the morphospecies praeedita which can be viewed as intermediate between eobulloides and edita. Eoglobigerina edita is an intermediate member of the eobulloides-edita-spiralis lineage which terminated in Zone P2 (see also Pearson, 1993).

Examination of the holotypes of Eoglobigerina hemisphaerica Morozova, E. theodosica Morozova, E. tetragona Morozova, and E. pentagona Morozova suggest that these are junior synonyms of edita Subbotina. These Morozova species are all characterized by having a weakly cancellate test and an elevated initial whorl which places them within the edita plexus. The difference between these morphotypes is the elevation of the spire (high in hemisphaerica and pentagona, lower in theodosica and tetragona) and the number of chambers in the ultimate whorl (5 1/2 in hemisphaerica, 5 in theodosica, 4 1/2 in pentagona, 4 in tetragona). The 5 1/2  chambered, finely cancellate holotype of hemisphaerica has a distinctly elevated spiral side; Morozova (1961) drew attention to the "stepped nature" of the antepenultimate and penultimate whorls which was said to serve to distinguish hemisphaerica from edita. We regard this as being within the range of variation of E. edita.

Stable isotopes: No data.

Stratigraphic range: Zone P to P1c; ?P2.

Global distribution: Worldwide in high and low latitudes.

Origin of species: This species evolved from Eoglobigerina eobulloides near the base of Zone P by an increase in the number of chambers in the last whorl and a slowing of the rate of growth of these chambers. In addition, there is a trend towards an elevated initial spire. An increase in the degree of spinosity is also evident in the evolution of E. edita.

Repository: Holotype No. 3975, St. Petersburg VNIGRI. Holotype examined by WAB.

Globigerina (Eoglobigerina) eobulloides Morozova, 1959, p. 1115, text-figs. 1a-c [Zone I - Zone of smooth-walled globigerinids (eoglobigerinids), lower Danian (Uylinian Stage), Tarkhankut Peninsula, Crimea, SW Russia].

Eoglobigerina eobulloides eobulloides (Morozova).-Blow, 1979, p. 1214, pl. 60, fig. 9 and pl. 61, fig. 1 (Zone P , DSDP Hole 47.2/11/4: 148-150 cm); pl. 65, figs. 8,9 and pl. 66, figs. 6,9 (Zone P , DSDP Hole 47.2/11/3: 148-150 cm); pl. 70, figs. 3,4 (Zone P1, DSDP Hole 47.2/11/3: 0-5 cm); pl. 73, fig. 6 (Zone P1, DSDP Hole 47.2/11/1: 148-150cm), Shatsky Rise, NW Pacific Ocean.

Eoglobigerina eobulloides (Morozova).-Huber, 1991c, p. 461, pl. 2, figs. 9-11 (Zone AP1A, ODP Hole 738C/20R/5: 376.54 mbsf; Kerguelen Plateau, Southern Indian Ocean).-Hemleben, M¸hlen, Olsson and Berggren, 1991, p. 126, pl 7, figs. 1-3 (Zone P , Clayton Basal Sands, Millers Ferry, Alabama).- Olsson, Hemleben, Berggren and Liu, 1992, p. 197, pl. 1, figs. 1-7 (Zone P , Clayton Basal Sands, Millers Ferry, Alabama).

Globigerina fringa Premoli Silva and Bolli, 1973 (non Subbotina, 1950), p. 541, pl. 7, figs. 6,9 (Zone P , DSDP Site 15/152/10/1: 127-130 cm; eastern Caribbean Sea)

Eoglobigerina fringa Smit, 1982 (non Subbotina, 1950), p. 329, pl. 2, figs. 11a-c (base of Zone P , Gredero section, SE Spain)

Eoglobigerina fringa Stott and Kennett, 1990, (non Subbotina, 1950), p. 559, pl. 1, figs. 5,6 (Zone AP , ODP Hole 690C/15X/4: 28-30 cm; Maud Rise, Weddell Sea, Antarctic Ocean).

Original description: Test with a flattish spiral; 4-4 1/2 chambers per whorl. Diameter 0.225 mm., height 0.13 mm. Differentiated from G. (Globigerina) bulloides d'Orbigny and G. (G.) pseudobulloides Plummer by its small aperture and smooth or definitely micro-cellular test wall.

Diagnostic characters: This spinose species has a moderately elevated trochospire, globular chambers, an umbilical to slightly extraumbilical rounded aperture which is bordered by a narrow, slightly flaring lip. Four to 4 1/2 chambers in the final whorl increase moderately in size. The umbilicus is small but open to the previous chambers. The cancellate wall texture is very weakly developed and difficult to view with the light microscope, especially where preservation of the wall is poor. The pores which are on average about 1 um in diameter at the narrowest point occur at the base of a shallow pit which is surrounded by the cancellate ridges. The wall ranges from 4 um to 7 um in thickness and the overall size of the test is generally less than 250 um.

Discussion: Considerable uncertainty and confusion surrounds the identification of this species and Globigerina fringa Subbotina, 1950 due to the very small size and generalized drawing of the holotype. Examination of the holotype (WAB and FR) under a light microscope shows it to be similar to E. eobulloides in general morphology and, thus, a possible senior synonynm. However, scanning electron micrographs (SEM) taken by FR of the two species show that they are distinctly different species. "Globigerina" fringa has a coarsely cancellate wall similar to that of Subbotina cancellata Blow, 1979. This type of cancellate wall texture is more advanced than that seen in Zone P globigerinids which suggests that fringa was described from a higher Danian stratigraphic level. See discussion under Subbotina cancellata for additional data on this species.

Four chambered forms of E. eobulloides were named by Blow (1979) as the subspecies E. eobulloides simplicissima. This form seems to be the same as Globigerina moskvini Shutskaya, 1953, a name that could be used for E. eobulloides except that some of the Shutskaya collections were destroyed during the 1960's and, thus the usage of this name is not advisable. Blow's (1979) illustrations of E. eobulloides agree well with the holotype. The small size of E. eobulloides and its very thin pore filled walls makes this species very susceptible to dissolution which is common in lowermost Danian sections.

Stable isotopes: Eoglobigerina eobulloides has a 13C and 18O signature similar to Parasubbotina and Globanomalina but typically with heavier 18O and more negative 13C than coexisting Praemurica and Woodringina (D'hondt and Zachos, 1993; Berggren and Norris, in press). The species shows little change in 18O or 13C over a large size range (Berggren and Norris, in press) although a positive size/ 13C relationship has been reported for the species at small shell sizes from the early Danian (D'Hondt and Zachos, 1993)

Stratigraphic range: Upper Zone P0 to P1b.

Global distribution: Worldwide in high to low latitudes.

Origin of species: Eoglobigerina eobulloides evolved from Hedbergella monmouthensis (Olsson) in late Biochron P0 by development of a spinose cancellate wall texture (Liu and Olsson, 1994). It is the earliest spinose taxon of the Cenozoic and, as such, it represents a completely new adaptive innovation (carnivory?) following the terminal Cretaceous event(s). Olsson et al. (1992) suggested that the spinose eobulloides lies at the base of an early Paleocene (Danian) radiation of normal perforate, cancellate spinose forms referable to Eoglobigerina and Subbotina (see also Pearson, 1993).

Repository: Holotype No. 3508/1, Moscow GAN. Examined by WAB and FR.

Globigerina spiralis Bolli, 1957a, p. 70, pl. 16, figs. 16-18 (Gt. uncinata Zone, Lower Lizard Springs Fm., Trinidad).-Bolli and Cita, 1960, p. 12, pl. 32, figs. 2a-c (Gt. trinidadensis-G. daubjergensis Zone, Paderno d'Adda section, N Italy).-Hillebrandt, 1962, p. 130, pl. 11, figs. 14, 15 (Zone E, lower Paleocene, Reichenhall-Salzburg Basin, Austro-German border).

Eoglobigerina spiralis (Bolli). -Blow, 1979, p. 1222, pl. 79, figs. 5-9 (Zone P2, DSDP Hole 20C/6/4: 72-74 cm, Brazil Basin, South Atlantic Ocean).

Igorina spiralis (Bolli). -Huber, 1991c, p. 461, pl. 3, figs. 13-15 (Zone AP2, ODP Hole 738C/16R: 338.50 mbsf, southern Kerguelen Plateau, Southern Ocean).

Original description: Shape of test medium to high trochospiral, biconvex, spiral side distinctly convex, umbilical side less so; equatorial periphery lobate; axial periphery rounded. Wall calcareous, perforate, surface smooth. Chambers inflated, globular or slightly compressed laterally; about 15, arranged in 3 whorls; the 5-6 chambers of the last whorl increase moderately in size. Sutures on spiral side radial or slightly curved, depressed; on umbilical side radial, depressed. Umbilicus narrow, open. Apertures distinct arches with faint lips, interiomarginal, umbilical; that of last chamber in some specimens tens to an extraumbilical-umbilical position. Coiling random. Largest diameter of holotype 0.28 mm.

Diagnostic characters: This spinose species has a distinct elevated spire and 4 1/2 to 5 1/2 chambers in the ultimate whorl. The chambers are closely appressed with an umbilically directed aperture. The radially directed intercameral sutures on the umbilical side appear as distinct valleys when surrounded by gametogenetic calcification. A thin discontinuous lip borders the aperture. The wall texture is more strongly developed than in the ancestral species, E. edita. The umbilicus is very small and is often overlapped by the ultimate chamber. The spine holes which are set along the cancellate ridges appear not to be as numerous as in E. edita.

Discussion: Bolli (1957a) regarded E. spiralis as the immediate ancestor of Igorina pusilla (Bolli). Although Blow (1979) agreed with Bolli, he noted that the transition from spiralis to pusilla involved the loss of a porticus as well as the extraumbilical extension of the aperture. Blow placed great emphasis on the importance of the porticus as distinct from an apertural lip. However, the study of Olsson et al. (1992) on wall texture of Danian globigerinids and globorotaliids showed that the separation of these two kinds of apertural apparatuses is erroneous as the apparent differences noted by Blow are due to ontogenetic and gametogenetic calcification. The fact that Blow regarded the spiralis-pusilla-albeari lineage as "so remarkably taxonomically and stratigraphically isolated" indicates that he considered the morphological change involved in the origin of pusilla (i.e. Igorina) as quite distinct from the mainstream globigerinacean evolutionary history. His belief holds true to some degree in regards to the origin of the Igorina lineage (see discussion under this genus). However, the ancestral relationship of spiralis to pusilla must be ruled out because the former species has a spinose wall texture whereas the latter has a nonspinose, praemuricate wall texture. Eoglobigerina spiralis is the end member of the eobulloides-edita-spiralis lineage which terminated in Zone P2 (see also Pearson, 1993).

Our work has revealed a spinose wall texture in E. spiralis and suggests that these spinose forms are unrelated to the non-spinose, igorinids. However, we have recognized an apparently nonspinose, wall texture in some specimens that we have initially attributed to E. spiralis. Indeed, the holotype of E. spiralis is too poorly preserved to verify the original presence or absence of spines. Consequentially, we are not completely sure of the generic designation of the species. Should paratypes of E. spiralis prove to have a non-spinose wall texture, we would be forced to consider this taxon as a possible ancestor to I. pusilla as originally suggested by bolli (1957a) and regard the spinose homeomorphs as derivatives of Eoglobigerina edita.

Stable isotopes: No data available.

Stratigraphic range: Zone P2; ?uppermost P1c.

Global distribution: Apparently worldwide in high to low latitudes.

Origin of species: Evolved from E. edita near the base of Zone P2 by the development of a tighter coil, a distinct elevated early coil of chambers, and a more strongly cancellate test wall.

Repository: Holotype (USNM P5030) deposited in the Cushman Collection, U.S. National Museum. Examined by WAB, BH, and RKO.

Type species: Globigerina pseudobulloides Plummer, 1926

Original description: Test very low trochospiral with 10-12 chambers, and with 4 to 5 chambers in the ultimate whorl. The chambers which are inflated globular and slightly ovoid in shape increase rapidly in size. The aperture is interiomarginal, umbilical to extraumbilical, a high rounded arch which is bordered by a narrow lip. The umbilicus is narrow, deep and open to the previous chambers. The wall is weakly to strongly cancellate and spinose. Spine holes are numerous and located at the juncture of and along the cancellate ridges. They may be obscured by gametogenetic and/or diagenetic calcification.

Discussion: The genus is distinguished by its very low trochospiral test, chambers that increase rapidly in size in the ultimate whorl, and by the high-arched umbilical to extraumbilical aperture. The number of chambers never exceeds 5 in the ultimate whorl. The last two chambers may be offset towards the umbilicus (P. variospira) giving the test the appearance of a higher spire.

Globigerina pseudo-bulloides Plummer, 1926, p. 133, pl. 8, figs. 9a-c (Zone P2, Wills Point Formation, Midway Group (upper Danian), Navarro County, Texas).

Globigerina pseudobulloides Plummer. -Subbotina, 1950, p. 106, pl. 4, figs. 8-10 (Danian Stage, N Caucasus). -Troelsen, 1957, p. 128, pl. 30, figs. 6a-c (Tylocidaris oedumi Zone, Hojerup, Stevns Klint), figs. 7a-c (basal Danian, Bogelund), figs. 8a-c (?Tylocidaris oedumi Zone, Hjerm) all Danian Stage, Denmark.-Bolli and Cita, 1960, p. 385, pl. 33, figs. 4a-c (Gt. trinidadensis-Gt. pseudobulloides Zone, Paderno d'Adda section, N Italy).-Gohrbandt, 1963, p. 44, pl. 1, figs. 7-9 (lower Paleocene, N of Salzburg, Austria).

Globorotalia pseudobulloides (Plummer). -Bolli, 1957a, p. 72, pl. 17, figs. 19-21 (Gt. pusilla pusilla Zone, lower Lizard Springs Fm., Trinidad).-Loeblich and Tappan, 1957a (partim), p. 192, pl. 40, figs. 3a-c (Tylocidaris oedumi Zone, Danskekalk Fm., Stevns Klint, Denmark, type Danian Stage); pl. 41, figs. 1a-c (Zone P1, McBryde Fm., Maryland); pl. 42, figs. 3a-c (Zone P1, Brightseat Fm., Maryland); pl. 43, figs. 3a-c (Zone P1, Kincaid Fm., Midway Group, NE Texas; non 4a-c = indeterminate abortive individual); pl. 44, figs. 6a-c (Wills Point Fm., Midway Group, NE Texas; non figs. 4,5 = indeterminate abortive individuals); pl. 45, figs. 1a-2c (Zone P3, Matthews Landing Marl Member, Porters Creek Clay, Wilcox County, Alabama); pl. 46, figs. 6a-c (Coal Bluff Marl Member, Naheola Fm., Wilcox County, Alabama); non pl. 40, figs. 9a-c (= Praemurica pseudoinconstans (Blow), paratype, Pine Barren Member, Clayton Fm., Wilcox County, Alabama).-Olsson, 1960, p. 46, pl. 9, figs. 19-21 (Zone P1, Hornerstown Fm., New Jersey)

Globorotalia (Globorotalia) pseudobulloides (Plummer). -Hillebrandt, 1962, p. 124, pl. 12, figs. 2a-c (lower Paleocene, Reichenhall-Salzburg Basin, Austro-German border).

Globorotalia (Turborotalia) pseudobulloides (Plummer).- Blow, 1979, p. 1096, pl. 69, figs. 2,3 (Zone P1, DSDP Hole 47.2/11/3:0-5 cm, Shatsky Rise, NW Pacific Ocean); pl. 71, figs. 4,5 (Zone P1, Hole 47.2/11/1: top section, Shatsky Rise, NW Pacific Ocean); pl. 75, figs. 2,3 (Zone P1, Lindi area, Tanzania, E Africa); pl. 248, figs. 6-8 (Zone P2, topotypes from Plummer locality 23, Navarro County, Texas); pl. 255, figs. 1-6 (Zone P1c?, Karlstrup, Denmark, upper Danian).

Subbotina pseudobulloides (Plummer).- Berggren, 1992, p. 563, pl. 1, figs. 7,8 (Zone P1b, ODP Hole 747A/19H/CC, Kerguelen Plateau, Southern Indian Ocean).

Parasubbotina pseudobulloides (Plummer). -Olsson, Hemleben, Berggren, and Liu, 1992, p. 197, pl. 3, figs. 1-7 (Figs. 1-5, Zone P1a, Pine Barren Member, Clayton Fm., Alabama), figs. 6,7 (Zone P2, upper Midway Fm., Texas).

Original description: Test rotaliform, very obtusely trochoid to plane dorsally, composed of about two and one-half convolutions, of which the last consists most generally of 5 (rarely 6) highly ventricose chambers increasing rapidly in size; periphery broadly rounded and lobate; shell wall thin and distinctly punctate but finely reticulate; superior face bearing a spire of small chambers only very slightly elevated, if at all, above the circumambient chambers of the final whorl; inferior face less convex and with a very distinct, though not large, umbilical depression; aperture a single, moderately large, lunate opening on the last chamber extending from the margin to the umbilicus and edged with a narrow, delicate, flaring lip.

Diameter up to .4 mm.

Diagnostic characters: Test very low trochospiral with 10-12 chambers, and with 5 chambers in the ultimate whorl. The inflated, globular chambers which are slightly ovoid in shape increase rapidly in size. The aperture is interiomarginal, umbilical to extraumbilical, a high rounded arch which is bordered by a narrow lip. The umbilicus is narrow, deep and open to the previous chambers. The cancellate spinose wall is weakly developed in the early forms of this species but becomes stronger in later forms. Spine holes are numerous and located at the juncture of and along the cancellate ridges. They may be obscured by gametogenetic and/or diagenetic calcification. The overall size of the test is generally greater than 250 um.

Discussion: The demonstration that P. pseudobulloides has a cancellate, spinose wall texture (Olsson et al., 1992) and is a member of a relatively minor offshoot of the early eoglobigerinid radiation, has given pause to long held notions about early Paleocene planktonic foraminiferal phylogenies (see also Pearson, 1993). This is because it has been commonly accepted for nearly 40 years that pseudobulloides was the ancestor of the post-Danian muricate morozovellid radiation (Bolli, 1957a; Blow, 1979). Parasubbotina pseudobulloides is a member of a lineage which includes P. varianta and apparently P. variospira. The lineage appears to have disappeared without descendants in the lower part of Zone P4.

The identification of pseudobulloides must be made with care because of the general, superficial similarity with Praemurica pseudoinconstans (see also Blow, 1979) in addition to the fundamental difference in wall texture between Parasubbotina and Praemurica which may not be evident in poorly preserved specimens.

Stable isotopes: Parasubbotina pseudobulloides has a positive 18O and negative 13C similar to Subbotina, Eoglobigerina, and Globanomalina. Both 18O and 13C display little size-related variability (DíHondt and Zachos, 1993; Berggren and Norris, in press).

Stratigraphic range: Uppermost Zone P to Zone P3a; ?P3b.

Global distribution: Worldwide in low to high latitudes.

Origin of species: Evolved from 4 1/2 to 5 chambered, weakly cancellate, spinose morphotypes identified as Parasubbotina sp. aff. pseudobulloides (Olsson et al., 1992, pl. 4, figs. 1-4) which occurs at the top of Zone P0 and in Zone P . They are not the same as the 5 to 6 chambered forms identified as G. (T.) aff. pseudobulloides by Blow (1979), which Blow regarded as having a phylogenetic relationship with pseudoinconstans.

Repository: Cotypes: Walker Museum Coll. 33076, Station 23

Parasubbotina aff. pseudobulloides (Plummer, 1926). -Olsson, Hemleben, Berggren, and Liu, 1992, p. 197, pl.4, figs. 1-4 (upper Zone P0 and Zone P , Millers Ferry, Alabama).

Diagnostic characters: A small, very low trochospiral test with 4 1/2 to 5 chambers in the ultimate whorl, globular chambers which increase rapidly in size, a high arched umbilical-extraumbilical aperture with a thin lip, and a weakly developed cancellate wall, spinose. Maximum diameter less than 225 um.

Discussion: This small morphotype apparently has often been identified as Globigerina fringa Subbotina, 1950. This species, however, has a coarsely cancellate wall and morphologic characters belonging to Subbotina. Parasubbotina aff. pseudobulloides is smaller than P. pseudobulloides and it has a very weakly developed cancellate wall in contrast to the strongly developed cancellate wall of pseudobulloides. The spinose condition is already present. As the cancellate wall becomes more strongly developed along with an increase in size this morphotype grades into pseudobulloides. Because of this fact, it is often difficult to consistantly identify the first occurrence of pseudobulloides and hence the base of Zone P1a. A more useful criterion for the base of Zone P1a is the last occurrence of Parvularugoglobigerina eugubina (Luterbacher and Premoli Silva).

Stable isotopes: there are no existing isotope data that are unambiguously attributed to this taxon. Data for P. pseudobulloides from the early Danian show that this taxon has 18O and 13C similar to Eoglobigerina and distinctly heavier 18O than Woodringina (DíHondt and Zachos, 1993).

Stratigraphic range: Upper part of Zone P0 to Zone P .

Global distribution: Probably widespread but few reliable identifications have been made.

Origin of species: Evolved from Hedbergella monmouthensis in Biochron P0 by a more rapid increase in chamber size and the development of a primitive cancellate wall texture and by becoming spinose.

Repository: Micropaleontology collections, University of Tuebingen, träger 9120-27; 9121-10. Examined by ChH and RKO.

Globigerina varianta Subbotina, 1953, (partim), p. 63, holotype: pl. 3, figs. 5a-c; paratypes: figs. 6a-7c, 10a-11c (Zone of rotaliiform Globorotalia, Elburgan Fm., Kuban River section, North Caucasus); figs. 12a-c (Zone of compressed Globorotalia, base of Lower White Series, Murzai-Tai, Mangyshlak Peninsula, SW Russia); pl. 4, figs. 1a-3c (Zone of rotaliiform Globorotalia, base Foraminiferal Layer F1, Khieu River section, near Nal'chik, North Caucasus).

Globorotalia (Globorotalia) varianta (Subbotina). -Hillebrandt, 1962, p. 125, pl. 12, figs. 10a-c, 11a,b (Zone D, Reichenhall-Salzburg Basin, Austro-German border).

Globorotalia (Turborotalia) quadrilocula Blow, 1979, p. 1109, holotype: pl. 87, fig. 7 (Zone P3, DSDP 47.2/10/1: 72-74 cm); paratypes: pl. 75, fig. 8 (Zone P1, Lindi, Tanzania), pl. 78, figs. 2-4 (Zone P2, DSDP 20C/6/4: 72-74 cm), pl. 83, fig. 3 (Zone P2, DSDP 47.2/10/3: 78-80 cm), pl. 87, fig. 8 (Zone P1, Lindi, Tanzania).

Subbotina varianta (Subbotina). -Berggren, 1992, p. 563, pl. 1, fig. 3 (ODP 747A/19H/CC, southern Kerguelen Plateau).

Original description: Shell globigerinelliform plano-convex, oval outline, consisting of 2 whorls with 4 chambers to the last whorl, rapidly increasing in size and closely packed together. The chambers to the last whorl, as is usual in the genus, are strongly inflated.

Dorsal side compressed, ventral convex. The chambers on the dorsal side are clearly defined and compressed whereas on the ventral side they are inflated and hemispherical. The last chamber is particularly large and is almost spherical. Sutures short, very slightly curved almost straight. On the dorsal side they are not deeply incised, on the central side they are strongly incised. Peripheral margin broadly rounded and coarsely scalloped. Umbilicus always distinct though small and having the form of a shallow depression in the middle of the ventral surface. Orifice slit-like, usually with small, indistinct lips forming a lamelliform border to the slit. Orifice along marginal suture. Dimensions: diameter 0.28-0.50 mm, thickness 0.12-0.24 mm.

Diagnostic characters: The species has a very low trochospiral test with generally four rapidly expanding chambers in the ultimate whorl. The wall is cancellate spinose with spine holes on the interpore ridges. The aperture is umbilical-extraumbilical, a rounded high arch bordered by a fairly broad continuous lip. The small, rounded umbilicus is deep and open to the surrounding chambers.

Discussion: The species is smaller and has fewer chambers in the ultimate whorl than does the closely related Parasubbotina pseudobulloides. Blow (1979) regarded P. varianta as an ecophenotypic variant of pseudobulloides comprising a rather specialized side branch of the pseudobulloides plexus. At the same time he erected the species Globorotalia (Turborotalia) quadrilocula which he considered to have originated from a "four-chambered last-whorl variant" of pseudobulloides. The holotype of varianta is a four-chambered form with an ultimate chamber somewhat more inflated than Blow's concept of quadrilocula. We regard quadrilocula as falling within the morphologic range of variability of varianta.

Stable isotopes: Parasubbotina varianta has a dC13C and 18O signature similar to P. pseudobulloides, Subbotina triloculinoides, and Globanomalina compressa but typically with heavier 18O and more negative 13C than coexisting Praemurica and Morozovella. The species shows little change in 18O or 13C over a large size range.

Stratigraphic range: Zone P1c to Zone P4 (basal part).

Global distribution: Worldwide in high and low latitudes.

Origin of species: Blow's observations on the close relationship of this species with pseudobulloides is supported by our observations. The species evolved in Zone P1c from pseudobulloides by a reduction in the number of chambers in the ultimate whorl and a tightening of the whorl.

Repository: Holotype No. 3994, paratypes No. 3995-4003, 4215, St. Petersburg VNIGRI (378/20). Examined by FR.

Globorotalia (Turborotalia )variospira Belford 1984, p. 18, pl. 24, figs. 15-17; pl. 25; figs. 1-7 (WABAG Sheet area, Papua, New Guinea).

Morozovella variospira (Belford), 1984, emended Van Ejden and Smit, 1992, p.113 , pl. 5, figs. 1-8; text-figs. 26: A-D ( Zone P3, ODP Hole 758A/30X/CC).

Original description: Test large, trochoid, consisting of about 12 chambers arranged in three whorls, usually 5 but sometimes 4 chambers visible from ventral side. Equatorial periphery lobate, axial periphery rounded. Chambers increasing only slowly in size, four chambers in penultimate whorl, coiling in most specimens then becoming looser and more open, generally with five chambers in final whorl, umbilicus wide, open, shallow. Occasional specimens with final chamber directed towards umbilicus, coiling also becoming tighter, 4-4 1/2 chambers visible from ventral side. Sutures on both dorsal and ventral sides narrow, deeply depressed, radial. Test wall perforate, with small pustules at umbilical margin of early chambers of last whorl, otherwise finely pitted. Aperture interiomarginal, umbilical-extraumbilical, low elongate, not clearly observed.

Diagnostic characters: Large ( 0.4 to 0.7 mm in diameter), low to (less frequent) moderately high, loosely coiled trochospiral test with strongly lobulate outline; any or all of the last 3-4 chambers with inwardly projecting umbilical "teeth", aperture an interiomarginal, umbilical-extraumbilical arch; loose coiling between chambers in the final whorl occasionally produces secondary spiral apertures with apertural lips, wall texture spinose and distinctly cancellate with large pore pits.

Discussion: Although described a decade ago, this species has been overlooked in studies of Paleocene planktonic foraminiferal faunas. This absence of attention is surprising since P. variospira is large and homeomorphic with the Neogene species Neogloboquardina dutertrei. Van Eijden and Smith (1992) drew attention to the distinct features of this large mid-Paleocene taxon, and noted its globoquadrinid-like umbilical "teeth" which is unique among Paleocene species. Parasubbotina variospira overlaps for a short stratigraphic interval with Globanomalina pseudomenardii and early members of the acarininid radiation (nitida, mckannai, subsphaerica ) within the lowermost part of Zone P4.

Stable isotopes:Parasubbotina variospira has a 13C and 18O signature similar to P. pseudobulloides, P. varianta, Subbotina triloculinoides, and Globanomalina compressa but typically with heavier 18O and more negative 13C than coexisting Morozovella.

Stratigraphic range: Zone P3a to P4 (lower part).

Global distribution: This species has been observed at (sub)tropical sites in the Atlantic, Indian and Pacific Oceans.

Origin of species: Evolved from P. varianta by development of a more loosely coiled test and inflated chambers. Both P. varianta and P. variospira share enlarged apertural flaps. These structures are restricted to the final chamber of P. varianta, but are found on the last 3-4 chambers in the final whorl of P. variospira.

Repository: Holotype (CPC 21919) and paratypes (CPC 21920-21922) deposited in Commonwealth Paleontological Collection, Bureau of Mineral Resources, Canberra, Australia.

Type species: Globigerina triloculinoides Plummer, 1926, emended

Diagnostic characters: Low trochospiral, tripartite test with 10-12 chambers, with 3-4 rapidly inflating, globular chambers in the ultimate whorl. The aperture is interiomarginal, umbilical to slightly extraumbilical in some species, a low arch. The apertural lip varies from narrow to a fairly broad and distinct apparatus that extends over the umbilicus. The umbilicus is small and nearly closed by the tight coiling. The wall is cancellate and spinose with the spines set at the juncture of the cancellate ridges with or without spine collars. The cancellate texture varies from weak to very strong and from moderate to very coarse or distinctly honeycombed.

Discussion: Olsson et al. (1992) showed that Subbotina was spinose. The concept of the genus followed here is similar to that of previous workers except that it is emended to include the spinose character.

Globigerina fringa Subbotina, 1953, p. 62, pl. 3, fig. 3 (Caucasus, Anapa, Azov-Black Sea Flysch, Pecten horizon, Danian).

Subbotina triangularis cancellata Blow, 1979, p. 1284, holotype: pl. 80, fig. 7 (Zone P2, DSDP Hole 20C/6/4, 72-74cm, South Atlantic Ocean); paratypes: pl. 80, figs. 2-6, 8,9 (same sample as holotype).

Diagnostic characters: A tightly coiled test with 3 1/2 to 4 chambers in the ultimate whorl, an umbilically directed aperture which is bordered by a broad, somewhat irregular lip, and a coarse cancellate wall on all chambers of the test. The test is compact, rounded in outline and slightly lobulate.

Discussion: Blow (1979) recognized the significance of the coarsely cancellate wall as a primary taxonomic character. The coarsely cancellate wall is a diagnostic feature of the cancellata-velascoensis lineage. Blow figured only umbilical views of his new taxon. Spiral views show a rounded, somewhat lobulate outline, with the ultimate chamber varying in size from slightly smaller to distinctly larger than the penultimate chamber, and the coarse cancellate wall of the earlier chambers. The axial periphery is broadly rounded.

In his discussion of this species Blow emphasized the apertural lip which he identified as a porticus. He believed that the porticus was a feature that was added as a later ontogenetic structure to the test. He based his belief on his study of the structure which in a SEM (1979, pl. 238, fig. 6) he interpreted as an infilling of pore-pits. At that time little was known about the ontogeny and gametogenesis of planktonic foraminifera. It is quite clear now that the lip forms as an extension of the primary chamber wall and that the cancellate texture develops throughout ontogeny and is enhanced during gametogenesis. The cancellate texture does not develop on the apertural lip which, apparently, led Blow to interpret it as a separate, late stage structure.

Blow named cancellata as a subspecies of Subbotina triangularis (White). However, this species has a distinctly different wall texture than cancellata. The recognition of a coarsely cancellate Subbotina lineage separates cancellata from triangularis and elevates it to species level.

Considerable confusion exists on the correct identification of Globigerina fringa Subbotina, 1950. The species is mostly identified in the lower Danian in Cretaceous/Paleogene boundary sections and is regarded as one of the earliest Danian species. The small size of the original figures of the holotype and its morphologic similarity to Eoglobigerina eobulloides Morozova, 1959 suggested that these species are synonymous (see Toumarkine and Luterbacher, 1985). SEMs taken by FR of the holotypes of the two species shows this not to be the case. The distinguishing characteristic of fringa is a coarsely cancellate wall texture which is similar to that in Subbotina cancellata Blow (1979). Furthermore, the coarsely cancellate wall texture observed in fringa is an advanced development in the evolution of wall texture in the Danian and is not present in Zone P stratigraphic levels. In fact, fringa was described from the "Pecten Horizon" which is apparently an upper Danian horizon in the Elburgan Formation in the northwest Caucasus.

Coarsely cancellate morphotypes Subbotina cancellata occur in Zone P1c at DSDP Site 356 in the Southern Atlantic Ocean. They appear quite similar to the SEM of the holotype of Globigerina fringa Subbotina (1950). These morphotypes are smaller than typical cancellata and have 4 to 4 1/2 chambers in the ultimate whorl. Although Subbotina described this species as having 4 chambers in the ultimate whorl, she chose a 4 1/2 chambered specimen as the holotype. Most specimens in a suite of adults at Site 356 are 4 chambered. Small immature specimens often have 4 1/2 chambers in the ultimate whorl and these specimens have an umbilical to a slightly extraumbilical aperture. In the adult specimens the inner whorl is composed of 4 1/2 chambers. This ontogenetic progression proceeds from an Eoglobigerina morphotype to a Subbotina morphotype and suggests that cancellata may have evolved from E. eobulloides. However, the strong cancellate wall texture is a distinctive characteristic of Subbotina. In general test morphology the cancellata morphotypes at Site 356 are similar to Subbotina trivialis Subbotina, 1953, suggesting that cancellata evolved from this species by enhancement of the cancellate wall.

The development of a coarsely cancellate wall begins a lineage which extends through S. cancellata to Subbotina velascoensis (Cushman), 1925. Although it is tempting to apply the name fringa to the Site 356 cancellata morphotypes we recommend that this not be done pending further study on the morphologic characteristics of this taxon and on the stratigraphy and phylogeny of the coarsely cancellate Subbotinas in Zone P1.

Thus, it is clear that identification of fringa and its taxonomy should be considered carefully when using literature data in interpretative studies. For example, Brinkhuis and Zachariasse (1988) included fringa (together with edita) in the monophyletic genus Parvularugoglobigerina Hofker, 1978, emended, which is characterized by having a microperforate, nonspinose wall texture. Similarly, specimens identified by Keller (1989a,b) as fringa (and indeed, other forms as cf. edita, hemisphaerica and taurica) from the basal Paleocene of El Kef (Tunisia) are microperforate, nonspinose forms referable to Parvularugoglobigerina.

Stable isotopes: No data available.

Stratigraphic range: Zone P1c to Zone P4, currently known range.

Global distribution: The full geographic extent of this species at this time is unknown. The species has been identified in North Atlantic (Site 549) and South Atlantic (Sites 20C, 356) DSDP drill sites.

Origin of species: Believed to have evolved from S. trivialis in the middle part of Zone P1, possible P1b. It is the first species of the coarsely cancellate Subbotina lineage. A reduction in the number of chambers in the ultimate whorl to 3 1/2, and an increase in test size characterizes Zones P2 to P4 morphotypes.

Repository: British Museum of Natural History, BP Cat. No. 41/31; paratypes: BP Cat. No. 41/9, 41/13-15, 41/18, 41/20, 41/22.

Diagnostic characters: Test small, very low trochospiral, consisting of 4 to 4 1/2 chambers in the ultimate whorl with a lobulate periphery. Chambers globular to nearly spherical in shape, coarsely cancellate with strongly developed interpore ridges, spinose. Sutures sharply depressed, radial on umbilical and spiral sides. The umbilicus is small and open, a deep depression. Aperture umbilical in adult specimens, slightly extraumbilical in immature specimens, a slightly asymmetrical low arch which is bordered by a broad wide lip that has an irregular edge. Maximum diameter of test ranges from 0.15 to 0.24 mm.

Discussion: Subbotina aff. cancellata occurs in Zone P1c at DSDP Site 356 in the Southern Atlantic Ocean. They appear quite similar to SEM of the holotype of Globigerina Fringa Subbotina (1950). Considerable confusion exists on the correct identification of fringa. It is mostly identified in the lower Danian in Cretaceous/Paleogene boundary sections and regard as one of the earliest Danian species. The small size of the original figures of the holotype and its morphologic similarity to Eoglobigerina eobulliodes Morozova, 1959 suggested that these species are synonymous (See Toumarkine and Luterbacher, 1985). Scanning electron micrographs taken by F. Rogel of the holotypes of the two species shows thus not to be the case. The distinguishing characteristic of S. fringa is the coarsely cancellate wall texture which is similar to that in Subbotina cancellata Blow (1979). Furthermore, the coarsely cancellate wall texture observed in S. fringa is an advanced development in the evolution of wall texture in the Danian and is not present in Zone P stratigraphic levels. Although Subbotina described this species as having 4 chambers in the ultimate whorl, she chose a 4 1/2 chambered specimen as the holotype. In aff. cancellata most specimens in a suite of adults are 4 chambered. Small immature specimens often have 4 1/2 chambers in the ultimate whorl and these specimens have an umbilical to a slightly extraumbilical aperture. In the adult specimens the inner whorl is composed of 4 1/2 chambers. The ontogenetic progression proceeds from an Eoglobigerina morphotype to a Subbotina morphotype and suggests that aff. cancellata may have evolved from E. eobulloides. However, the strong cancellate wall texture is a distinctive characteristic of Subbotina. In general test morphology aff. cancellata is similar to Subbotina trivialis Subbotina, 1953, suggesting that it evolved from this species by enhancement of the cancellate wall. The development of a coarsely cancellate wall begins a lineage which extends through S. cancellata and Subbotina velascoensis (Cushman), 1925. Although it is tempting to apply the name fringa to the aff. cancellata forms we recommend that this not be done pending further study on the morphologic characteristics of this taxon and on the stratigraphy and phylogeny of the coarsely cancellate Subbotinas in Zone P1.

Thus, it is clear that identification of fringa and its taxonomy should be considered carefully when using literature data in interpretive studies. For example, Brinkhuis and Zachariasse (1988) included fringa (together with edita) in the monophyletic genus Parvularugoglobigerina Hofker, 1978, emended, which is characterized by having a microperforate, nonspinose wall texture. Similarly, specimens identified by Keller (1989a,b) as fringa (and indeed, other forms as cf. edita, hemisphaerica and taurica) from the basal Paleocene of El Kef (Tunisia) are microperforate, nonspinose forms referable to Parvularugoglobigerina.

Stratigraphic range: lower Danian. The exact stratigraphic range of this taxon is yet to be established. At DSDP Site 356 it occurs in Zone P1c and gives rise to cancellata near the base of Zone P2. It dos not occur in Zone P nor has it been observed so far in Zone P1a.

Global distribution: South Atlantic Ocean (Site 356). The morphologic characteristics of the coarsely cancellate Subbotinas suggest that they have a broad global distribution.

Origin of species: Believed to have evolved from S. trivialis in the middle par of Zone P1, possible P1b. It is the first species of the coarsely cancellate Subbotina lineage and is ancestral to S. cancellata.

Repository: Micropaleontology collections, University of Tuebingen, trager Ols 7.
 
 

Globigerina triangularis White, 1928, p. 195, pl. 28, figs. 1a-c (Paleocene, Velasco Fm., Mexico).-Bolli, 1957a, p. 71, pl. 15, figs. 12-14 (Globorotalia pseudomenardii Zone, lower Lizard Springs Fm., Trinidad).-Shutskaya, 1970a, p. 104, pl. 3, figs 5a-c (lower part Ac. tadjikistanensis djanensisZone, Khieu River section, Nal'ckik, N. Caucasus.-Shutskaya, 1970b, (partim) p. 118, pl. 20, figs 7a-c (Gt. angulata Zone, Chaaldzhin Group, Malyi Bakhan Ridge, W. Turkmenia); p. 220, pl. 23, figs 1a-c (lower part of Ac. tadjikistanensis djanensis Zone, Khieu River section, Nal'ckik, N. Caucasus); p. 224, pl. 25, figs. 1a-c (upper part of Ac. tadjikistanensis djanensis Zone, Khieu River section, Nal'chik, N Caucasus) non pl. 17, figs. 14a-c (= S. triloculinoides).

Globigerina triloculinoides Loeblich and Tappan, 1957a (partim) (non Plummer, 1926), p. 183, pl. 62, figs. 3a-c (Zone P4, Velasco Fm., Mexico); not pl.62, figs. 4a-c and other illustrations of S. triloculinoides.

Globigerina inaequispira Loeblich and Tappan, 1957a, p. 181, pl. 52, figs. 1a-2c (Zone P4, Vincentown Fm., New Jersey).

Subbotina patagonica/triangularis group Tjalsma, 1977, p. 510, pl. 4, fig. 2 (Zone P5, DSDP Site 329/32/4:67-68cm), figs. 3-6 (Zone P6, DSDP Site 329/4:139-141cm), SW flank of Falkland Plateau, SW Atlantic Ocean.

Subbotina triangularis triangularis (White). -Blow, 1979, p. 1281, pl. 91, figs. 7,9 (Zone P4, DSDP Hole 21A/3/6:74-76cm, South Atlantic Ocean); pl. 98, fig. 6 (Zone P4, given as P5, Lindi area, Tanzania, E. Africa); pl. 107, figs. 8,9 (Zone P4, given as P6, DSDP Hole 20C/6/3:76-78cm, South Atlantic Ocean).

Globigerina pseudotriloba Shutskaya, 1970a (non White, 1928), p. 85, pl. 2, figs. 7a-c (Ac. conicotruncana Zone, Urukh River section, N Ossetiya, N. Caucasus).

Globigerina uruchaensis Shutskaya, 1970a, p. 87, pl. 2, figs. 6a-c (holotype, Ac. conicotruncana Zone, Urukh River section, N Ossetiya, N. Caucasus).

Globigerina gerpegensis Shutskaya, 1970a, p. 104, pl. 3, figs. 3a-c (holotype, upper part of Ac. tadjikistanensis djanensis Zone, Kachan Stage, Lower Danatin Subgroup, Malyi Balkhan Ridge, W. Turkmenia).

Original description: Test triangular, composed of about two coils arranged in a low trochoid spire; chambers inflated, somewhat appressed, three sub-equal chambers comprising the last whorl; sutures deep; aperture must be in the small umbilicus.

Diameter of type specimen, 0.4 mm.; thickness, 0.2 mm.

Diagnostic characters: Test a somewhat loose coil of 3 1/2 chambers in the ultimate whorl, with a narrow, deep umbilicus which is sometimes obscured by an overlapping ultimate chamber. Shape of test triangular in umbilical view, the ultimate chamber often smaller than the penultimate one, low oval in shape. Axial periphery broadly rounded. Aperture umbilical to slightly extraumbilical in position, bordered by a thin, sometimes irregular lip. The wall is cancellate, spinose with an asymmetricallydeveloped pore pattern and well developed coalescing spine collars.

Discussion: Shutskaya's (1970a,b) references to pseudotriloba, triangularis, uruch (a) ensis and gerpegensis are all considered synonymous with Subbotina triangularis. Her criteria provided to distinguish between these forms and her text-illustrations are insufficient for consistent discrimination and, indeed, she (1970a, p. 104) included Bolli's illustration of triangularis (1957a, pl. 15, figs. 12-14 in the synonym of her new taxon, G. gerpegensis.

The distinctive spinose wall texture of Subbotina triangularis sets it apart from the trivialis-triloculinoides lineage and the cancellata-velascoensis lineage. Subbotina triangularis may be a stem form for a separate lineage that links with Eocene species but this possibility has not been investigated. It may possibly be linked with Globigerina praebulloides which has a similar wall texture.

Stable isotopes: Subbotina triangularis displays a more positive 18O and more negative 13C than coexisting Morozovella and Acarinina(D'hondt et al. 1994). The species shows little change in 18O or 13C over a large size range (D'Hondt et al. 1994).

Stratigraphic range: Zone P2 to Zone P5, ?P6.

Global distribution: Apparently has a global distribution in low to mid-latitudes.

Origin of species: Probably evolved from S. triloculinoides in Zone P3 by developing a more evolute coil, by an increase in test size, and a more asymmetical pore pattern with well developed coalescing spine collars.

Repository: Columbia University Paleontology Coll. No. 19881 from locality No. 11.

Globigerina triloculinoides Plummer, 1926, p. 134, pl. 8, figs. 10a-b (Zone P2, Wills Point Fm., Midway Group, Navarro Co., NE Texas, Station 23).-Bolli, 1957a, (partim), p. 70, pl. 15, figs. 18-20 (Gt. pusilla pusilla Zone, Lizard Springs Fm., Trinidad), non pl. 17, figs. 25, 26.-Loeblich and Tappan, 1957a, (partim), p. 183, pl. 40, figs. 4a-c (Tyl. oedumi Zone, Danian Stage, W. Denmark); pl. 41, figs. 2a-c (Zone P1c, McBryde limestone of Clayton Fm., Wilcox County, Alabama); pl. 42, figs. 2a-c (Zone P1c, Brightseat Fm., Maryland); pl. 43, figs. 5a-c (Zone P1, Kincaid Fm., Travis County, south Texas), figs. 9a-c (Zone P2, Wills Point Fm., Midway Group, Navarro County, NE Texas); pl. 45, figs. 3a-c (Zone P3, Matthews Landing Marl, Naheola Landing , Tombigbee River, Alabama); pl. 49, figs. 2a-c (Zone P3b, Hornerstown Fm., New Jersey) (non pl. 46, figs. 1a-c; pl. 47, figs. 2a-c; pl. 52, figs. 3-7; pl. 56, figs. 8a-c; pl. 62, figs 3,4) .-Bolli and Cita, 1960, p. 13, pl. 31, figs. 1a-c (Gt. trinidadensis-Gl. daubjergensis Zone, Paderno d'Adda section, N Italy).-Berggren, 1962, p. 86, pl. 14, figs. 1a-2b (Zone P1b, type Danian, Stevns Klint, Denmark) .-Hillebrandt, 1962, p. 119, pl. 11, figs 1a-c (Danian Stage, Richenhall-Salzburg Basin, Austro-German border). -Shutskaya, 1970b, p. 118, pl. 18, figs. 1a-c (upper subzone of Gl. trivialis-Gl. daubjergensis-Gl. compressa Zone, Malyi Balkhan Ridge, W. Turkmenia); pl. 19, figs. 3a-c (Ac. inconstans Zone, Malyi Balkhan Ridge, W. Turkmenia); pl. 21, figs. 5a-s (Ac. praepentacamerata Zone, Urukh River section, N Caucasus); pl. 23, figs. 12a-c (lower subzone of Ac. tadjikistanensis djanensis Zone, Malyi Bakhan Ridge, W. Turkmenia).

Subbotina triloculinoides (Plummer). - Brotzen and Po aryska, 1961, p. 160, pl. 4, fig. 4; p. 160, text-fig. 2 (lower Paleocene, Midway Group, Texas).-Belford, 1967, p. 7, pl. 1, figs. 1-5 (Paleocene, Australia). -Stott and Kennett, 1990, p. 559, pl. 2, fig. 12 (Zone AP1a, ODP Hole 690C/15X/2: 46-50 cm, Maud Rise, Weddell Sea, Antarctic Ocean).

Subbotina triloculinoides triloculinoides (Plummer). Blow, 1979, p. 1287, pl.74, fig. 6 (Zone P1, DSDP Hole 47.2/11/1:148-150 cm, Shatsky Rise, NW Pacific Ocean); pl. 80, fig. 1 (Zone P2, DSDP Hole 20C/6/4:72-74 cm, South Atlantic Ocean); pl. 98, fig. 7 (Zone P4, given as P5, Lindi area, Tanzania, East Africa); pl. 238, fig. 5 (same specimen as pl. 80, fig. r, q.v.); pl 248, figs. 9, 10 (topotypes, Zone P2, Navarro County, Texas); pl. 255, fig. 9 (Zone D2 of Bang, 1969, probably Zone P1c, Karlstrup, Denmark); pl. 257, fig. 9 (Zone P1c, upper Danian, Klagshamn, Sweden).

Globigerina pseudotriloba White, 1928, p. 194, pl. 27, figs. 17a,b (Velasco Fm., Mexico).

Globigerina stainforthi BrÃnnimann, 1952, p. 171, pl. 3, figs. 10-12 (Paleocene, Trinidad).

Globigerina (Globigerina) microcellulosa Morozova, 1961, p. 14, pl. 1, fig. 11 (Danian, Crimea, Tarkankut).

Original description: Test spiral, trochoid, composed of about 2 convolutions, the last of which is composed of 3 1/2 very rapidly increasing and highly globose chambers; periphery very broadly rounded and distinctly lobate; shell surface strongly reticulate; superior face rounded with a very low spire of neatly coiled tiny chambers of the preceding whorl; inferior face rounded with a very shallow umbilical depression; aperture a small arched slit on the last chamber and edged with a more or less prominent, delicately notched flap that extends from a point near the periphery to the umbilical depression.

Greatest diameter up to .35 mm; usually less.

Diagnostic characters: A medium sized, lobulate, trilobate test with 3-3 1/2 chambers in the ultimate whorl which increase moderately in size, the ultimate chamber occupies up to 1/2 of the test size. Intercameral sutures depressed, straight to slightly curved on umbilical and spiral sides of the test. Umbilicus narrow, deep, often covered by a well developed apertural lip. Aperture umbilical, slightly asymmetrical towards an extraumbilical direction. The test walls are strongly cancellate, spinose, the spines occurring at the juncture of and along the cancellate ridges.

Discussion: Together with Parasubbotina pseudobulloides (Plummer), this taxon is probably one of the most cited, yet most frequently misidentified taxa of the Paleocene. The detailed analysis of Blow (1979) may serve as a guide to a reasonably consistent delineation of the main characteristics of this species. Although S. triloculinoides has a strongly cancellate wall texture, its does not possess the symmetrical coarsely cancellate wall of the S. cancellata- S. velascoensis lineage. The ancestral S. trivialis is more weakly cancellate and S. triangularis has an asymmetrical cancellate surface with distinct spine collars. Unlike Blow (1979) we do not recognize the two subgroups (aside from S. triloculinoides s.s.) to which some sort of formal nomenclature may be applied: S. triloculinoides stainforthi (Bronniman) and S. triloculinoides nana (Khalilov). The former is interpreted as a junior synonym of S. triloculinoides, as has been more or less suggested by Bolli (1957a), and the "taxon" nana Khalilov is a squat, subquadrate form with a linaperta-like aperture referable to S. velascoensis (Cushman) and not to triloculinoides, (verified by examination of holotype by WAB). While Khalilov (1967) indicated that the range of nana is upper Paleocene to lower Eocene in the Malyi Caucasus (Azerbaizhan) and Maly Balkhan (Turkmenia), the morphology of velascoensis (=nana) appears already in Zone P3. The SEMs of the holotype of S. microcellulosa (Morozova) show this taxon to be a junior synonym of triloculinoides.

Stable isotopes: Subbotina triloculinoides has a 13C and 18O signature similar to P. pseudobulloides, P. varianta, G. compressa and G. planocompressa but typically with heavier 18O and more negative 13C than coexisting Praemurica and Morozovella. The species shows little change in 18O or 13C over a large size range (Berggren and Norris, in press).

Stratigraphic range: Zones P1b to P4. While its upper stratigraphic limit remains somewhat equivocal, we have not observed it above Zone P4 and believe it is restricted to the Paleocene.

Global distribution: Essentially worldwide distribution in low to high latitudes

Origin of species: While Blow (1979) was of the opinion that the origin of triloculinoides lay in the plexus of forms which radiate from Eoglobigerina eobulloides s.l. towards E. trivialis and S. triloculinoides and, more specifically, with E. eobulloides simplicissima, we view its evolution from trivialis to be a more likely solution (see also Olsson et al., 1992).

Repository: Walker Museum Collection 33076, Station 23. Holotype examined by WAB.

Globigerina trivialis Subbotina, 1953 (partim), p. 64, holotype: Pl. 4, figs. 4a-c; paratypes: figs. 5a-7c (Zone of rotaliiform Globorotalia (=P1), basal Elburgan Fm., Kuban River section, N. Caucasus); non pl. 4, figs. 8a-c (same horizon; = S. triloculinoides). -Shutskaya, 1970b, (partim), p. 118, pl. 18, figs. 2a-c (upper subzone of Gl. trivialis-Gl. daubjergensis-Gl. compressa Zone, Malyi Balkhan Ridge, W. Turkmenia); pl. 18, figs. 16a-c (middle subzone of Gl. trivialis-Gl. daubjergensis-Gl. compressa Zone, Malyi Balkhan Ridge, W. Turkmenia); non pl. 21, figs. 3a-c, pl. 23, figs. 6a-c.

Eoglobigerina trivialis (Subbotina).- Blow, 1979, p. 1224, pl. 65, figs. 1-3, pl. 66, figs. 4,7 (Zone P , DSDP Hole 47.2/11/3:148-150 cm, Shatsky Rise, NW Pacific Ocean); pl. 69, fig. 9, pl. 70, fig. 8 (Zone P1, DSDP Hole 47.2/11/3:0-5 cm); pl.70, figs. 1,2 (Zone P2, DSDP Hole 20C/6/4:72-74 cm, South Atlantic Ocean). -Stott and Kennett, 1991, p. 559, pl. 2, fig. 11 (Zone AP1, ODP Hole 690C/15X/2:46-50 cm, Maud Rise, Weddell Sea, Antarctic Ocean). -Berggren, 1992, p. 563, pl.1, fig. 1, ODP Hole 747C/19H/CC, Kerguelen Plateau, Southern Indian Ocean).

Eoglobigerina cf. trivialis (Subbotina). -Blow, 1979, p. 1229, pl. 55, fig. 9 (Zone P , DSDP Hole 47.2/11/5:148-150 cm, Shatsky Rise, NW Pacific Ocean).

Eoglobigerina aff. trivialis (Subbotina). -Blow, 1979, p. 1228, pl. 61,fig. 8 (Zone P , DSDP Hole 47.2/11/4:148-150 cm), pl. 69, fig. 8 (Zone P1, DSDP 6/47.2/11/3:0-5 cm), pl. 74, figs. 7,8 (Zone P1, DSDP 6/47.2/11/1:148-150 cm), Shatsky Rise, NW Pacific Ocean.

Subbotina trivialis (Subbotina). -Huber, 1991c, p. 461, pl. 3, fig. 16, 17 (Zone AP1, ODP Hole 738C/17R: 350.45 mbsf, Kerguelen Plateau, Southern Indian Ocean). -Olsson, Hemleben, Berggren and Liu, 1992, p. 202, pl. 4, figs. 5-8 (Zone P1a, Pine Barren Member, Clayton Fm., Millers Ferry, Alabama).

Original description: Shell inflated with tall first whorls which are 2 in number and large in size. The last whorl consists of 4-4 1/2 almost regular, spherical chambers which differ very slightly in size. The chambers are closely packed together and partially overlap each other. Because of this the shell is very compact. Peripheral margin broadly undulate; septal sutures curved, deeply incised as is characteristic of species with strongly inflated shells. Orifice small, and forming a slightly curved or straight slit which is situated along the marginal suture and near to, or more rarely above, the umbilicus. Walls with relatively large pores and a cellular pattern. Mean dimensions: diameter 0.40 mm, thickness 0.23 mm.

Diagnostic characters: A low trochospiral, tightly coiled test with 3 1/2 chambers in the ultimate whorl and an umbilical aperture with a thin lip. The ultimate chamber is equal to, or slightly smaller than the penultimate one. The wall is weakly cancellate and spinose. The spines are set at the junctures of the cancellate ridges. The umbilicus is small and nearly closed by the tight coiling. The overall size of the test is generally less than 250 um.

Discussion: Although Subbotinaís original description refers to 4 to 4 1/2 chambers in the final whorl, her illustration of the holotype and paratypes show only 3 1/2 chambers. Blow (1979) presented a thorough analysis of this morphospecies with which we in large part agree. For instance, like Blow we have not observed individuals as large as 0.4 mm in diameter as noted by Subbotina (1953) for her holotype from the N Caucasus. In fact, remeasurement of her holotype by FR shows a diameter of 0.35 mm. Blow also described forms with more closely appressed chambers and abortively developed final chambers (as Eoglobigerina cf. trivialis) and forms with a turreted, high-spired umbilical side (as Eoglobigerina aff. trivialis). We include these morphotypes here in the concept of trivialis, as they do not appear to have any significant numerical representation or stratigraphic continuity in occurrence.

Stable isotopes: No data available.

Stratigraphic range: Zone P to Zone P2.

Global distribution: Essentially worldwide in low to high latitudes.

Origin of species: While agreeing with Blow (1979) that the distinction between the wall texture of eoglobigerinids and subbotinids is one of degree, rather than kind, we view trivialis as the earliest representative of Subbotina (in contrast to Blow, 1979, who retained it in Eoglobigerina) and the main lineage that ultimately gave rise to the late Paleogene-Neogene spinose globigerinid radiation (see also Pearson, 1993; Liu and Olsson, 1994)

Repository: Holotype No. 4004, paratypes No. 4005-7 St. Petersburg VNIGRI. Holotype examined by FR.

Globigerina velascoensis Cushman, 1925, p. 19, pl. 3, figs. 6a-c (Paleocene, Velasco Fm., Tamalte Arroyo, Hacienda el Limon, San Luis Potosi, Mexico). -White, 1928, p. 196, pl. 28, figs. 2a-b (Paleocene, Velasco Shale, Tampico Embayment, eastern Mexico). -Bolli, 1957a, p. 71, pl. 15, figs. 9-11 (Zone P4, Lizard Springs Fm., Trinidad).-Bolli and Cita, 1960, p 374, pl. 34, figs. 8a-c (Gt. pseudomenardii Zone, Paderno d'Adda section, N Italy). -Hillebrandt, 1962, p. 120, pl. 11, figs. 4a-b (Paleocene, Reichenhall-Salzburg Basin, Austro-German border). -Gohrbandt, 1963, p. 47, pl. 2, figs. 1-3 (upper Paleocene, N of Salzburg, Austria). -Shutskaya, 1970a, p. 94, pl. 4, figs 3a-4c, 6a-c (figs. 3,6: Ac subsphaerica Zone, Kambileevka River, Black Mountains, N Caucasus; fig. 4: Ac. subsphaerica Zone, Tarkhankut Peninsula, Crimea).

Globigerina velascoensis Cushman var. compressa White, 1928, p. 196, pl. 28, figs. 3a,b (Velasco Fm., Tampico Embayment, eastern Mexico).

Globigerina triloculinoides Plummer nana Khalilov, 1956, p. 236, pl. 1, figs. 4a-c (upper Paleocene, holotype from Akhchakuima, NE foothills of Malyi Caucasus, Nakhichevan Autonomous Republic, Azerbaizhan).

Globigerina quadritriloculinoides Khalilov, 1956, p. 237, pl. 1, figs. 5a-c (upper Paleocene, Akhchakuima, NE foothills of Malyi Caucasus, Nakhichevan Autonomous Republic, Azerbaizhan).

Original description: Test much compressed, the dorsal side with all the chambers visible, ventral side only those of the last-formed coil, sides nearly parallel, periphery broadly rounded; chambers distinct, three or four making up the last-formed coil, early chambers subglobular, later ones becoming more compressed, and the inner margin fairly straight; wall finely and evenly reticulate; aperture on the ventral side, elongate.

Diameter 0.45 mm.; thickness 0.25 mm.

Diagnostic characters: A tightly coiled test with compressed chambers and a subquadrate test shape with an umbilical aperture. The ultimate chamber is much compressed, an elongate oval shape that makes up about half of the test and typically overhangs the earlier chambers. The aperture is bordered by a thin elongate lip that does not extend along the entire length of the aperture and is squared off where it terminates. The test wall is coarsely and symmetrically cancellate, spinose.

Discussion: Although the holotype specimen is deformed, Bolli (1957a) defined the concept of this species which is here followed. The species was described from the Velasco Shale from which White (1928) illustrated better preserved specimens which he identified with Cushman's species. White's concept was followed by Bolli. White's figures, although drawings, illustrate a quadrate shaped test with much compressed chambers. Furthermore, his drawings clearly indicate a symmetrical, coarsely cancellate test wall. Cushman (1925) in his description referred to an evenly reticulate wall which suggests that he observed the wall texture which characterizes this species.

Bolli (1957a) and Blow (1979) considered that S. velascoensis evolved from S. triangularis at, or near, the Zone P3/P4 boundary and believed that velascoensis represented a dead-end lineage. Subbotina velascoensis has a much different wall texture than does triangularis. The latter species has a finer asymmetrical cancellate wall with coalescing spine collars whereas velascoensis has a symmetrical coarsely cancellate wall texture. In velascoensis, the spines are set at the intersection of the cancellate ridges and are not supported by spine collars as in triangularis. We consider velascoensis as the end member of the symmetrical coarsely cancellate lineage that begins with S. cancellata. Subbotina cancellata, gives rise to velascoensis at, or near, the Zone P3/P4 boundary. We agree with Bolli and Blow that velascoensis represents a dead-end lineage.

Stable isotopes:Subbotina velascoensis has a 13C and 18O signature similar to P. varianta, G. compressa , and G. pseudomenardii but typically with heavier 18O and more negative 13C than coexisting Acarinina and Morozovella (Berggren and Norris, in press).

Stratigraphic range: Zone P3b to Zone P6a.

Global distribution: An essentially worldwide distribution in low to mid-latitudes with preferential development in the mid-higher latitudes as is common with the subbotinids.

Origin of species: Evolved from S. cancellata by the development of a more tightly coiled, quadrate test and compressed chambers.

Respository: U.S. National Museum, Cushman Collection No. 4348USNM. Examined by WAB and RKO.

Diagnostic characters: Test trochospiral with an umbilical to extraumbilical aperture with a prominent lip. Apertures of earlier formed chambers remain visible around the umbilicus.

Type species: Anomalina lorneiana d'Orbigny var. trochoidea Gandolfi, 1942

Diagnostic characters: Test low to very low trochospiral with with 18 or more globular chambers. The number of chambers in the ultimate whorl varies from 5-8. The aperture is interiomarginal, umbilical-extraumbilical, a low to high arch, bordered by a narrow lip. The umbilicus is small, deep and open to the apertures of previous chambers. The wall is normal perforate, smooth to heavily pustulose. Nonspinose.

Discussion: The late Maastrichtian species Hedbergella holmdelensis and H. monmouthensis show the reduced number of chambers in the ultimate whorl and less pustulose chambers. Late Cretaceous species of Hedbergella are generally typified by more numerous chambers in the ultimate whorl and by more pustulose chambers.

Hedbergella holmdelensis Olsson, 1964, p. 160, pl. 1, holotype: figs. 2a-c; paratype: figs. 1a-c (upper Maastrichtian, Navesink Fm., New Jersey). -Huber, 1994, p. 18-22, pl. 2, figs. 1-8 (upper Maastrichtian, Navesink Fm., New Jersey).

Original description: Test very low trochospiral, nearly planispiral in appearance, much compressed; equatorial periphery strongly lobate; axial periphery rounded. Wall finely perforate, delicately hispid. Chambers five, occasionally six, in final whorl, increasing rather rapidly in size, moderately inflated, ovate, elongate in direction of coiling compressed ovate in axial view. Sutures depressed and gently curved on spiral side, depressed and radial on umbilical side. Aperture a low arch, interiomarginal, umbilical-extraumbilical, bordered by a thin lip; apertures of previously formed chambers open into the umbilicus.

Greatest diameter of holotype 0.20 mm., greatest thickness 0.10 mm. Unfigured paratypes range from 0.15 to 0.24 mm. in maximum diameter, from 0.07 to 0.12 mm. in maximum thickness. Greatest diameter of figured paratype 0.26 mm., greatest thickness 0.10 mm.

Diagnostic characters: This species is characterized by the small, nearly planispiral test with 5, occasionally 6 chambers in the ultimate whorl. Six to seven chambers make up the penultimate whorl. The ovate, elongated, compressed chambers are typical of the species. The axial periphery is imperforate for the most part, being only sparsely penetrated by pores. The wall surface is smooth but is lightly covered by small pustules. The aperture, a high arch that extends from the umbilicus to the axial periphery, is bordered by a thin, narrow, uniformly wide lip.

Discussion: Hedbergella holmdelensis is the ancestral species of the genus Globanomalina which first appears in Zone P0 of the basal Danian. Morphologically, H. holmdelensis is very similar to the first species of Globanomalina, G. archeocompressa. The key morphologic characters that phylogenetically link the two species are the small, compressed smooth-walled test, the imperforate axial periphery, and the umbilical-extraumbilical aperture with its narrow lip. Hedbergella holmdelensis retains two typical hedbergellid characters, however, which are; the numerous, spherical-shaped chambers in the inner whorl, an adult character of the majority of hedbergellid species in the Cretaceous and the pustulose surface of the test which is distributed rather uniformly over the chambers, in contrast to the more restricted occurrence of pustules in Paleogene species of Globanomalina. Pustule growth, particularly in the umbilical area, is related to gametogenetic calcification in smooth-walled species in the Cenozoic. Pustule growth in Hedbergella is largely unexplored and its relationship to gametogenesis is unknown. Nevertheless, the difference in its distribution on the test suggests a different biological function for this genus than for Globanomalina. Globanomalina, however, is placed in the Hedbergellidae to reflect its phylogenetic linkage with this important Cretaceous group of planktonic foraminifera.

Stable isotopes: No data available

Stratigraphic range: Lower Maastrichtian to lower Zone P0.

Global distribution: Apparently worldwide in high to low latitudes.

Origin of species: Evolved in the Maastrichtian from a multichambered ancestor.

Repository: Holotype (USNM 640917) and paratype (USNM 640919) deposited in the Cushman Collection, U.S. National Museum. Examined by BTH and RKO.

Globorotalia monmouthensis Olsson, 1960, p. 47, pl. 9, figs. 22-24 (upper Maastrichtian, Redbank Fm., New Jersey)

Hedbergella monmouthensis (Olsson). -Olsson, 1964, p. 160, pl. 1, figs. 3a-c (upper Maastrichtian, Redbank Formation, New Jersey). -Sliter, 1976, p. 542, pl. 3, figs. 1-3 (Maastrichtian, DSDP Hole 327A/12/1: 69-72 cm, eastern Falkland Plateau, South Atlantic Ocean). -Huber, 1990, p. 503, pl. 2, figs. 6-8, pl. 6, fig. 2 (Maastrichtian, ODP Hole 690C/20X/5: 108-110 cm, Maud Rise, Southern Ocean). -Huber, 1991a, p. 292, pl.1, figs. 2, 3 (Abathomphalus mayaroensis Zone, ODP Hole 698A16R/2: 67-71 cm, Northeast Georgia Rise, Southern South Atlantic Ocean). -Huber, 1994, p. 22-25, pl. 2, figs. 9-16 (upper Maastrichtian, Redbank Fm., New Jersey).

Original description: Test.--very low trochospiral, nearly planispiral in appearance. Equatorial periphery strongly lobate. Axial periphery rounded. Wall.--calcareous, finely perforate, covered with short minute spines. Chambers.--inflated, globular; about 15 arranged in 2 1/2 whorls. The 5, occasionally 6, chambers of the final whorl increase rather rapidly in size; the last two make up over 1/2 of the test. Sutures.--spiral side: oblique in early stages to straight in adult stages, depressed, Umbilical side: radial, depressed. Umbilicus.--narrow, deep. Aperture.--a low arch with a distinct lip; interiomarginal, extraumbilical-umbilical. Coiling.--random. Diameter 0.20 to 0.25 mm.

Diagnostic characters: This small species has 5 to 5 1/2 inflated chambers in the ultimate whorl that rapidly increase in size. The penultimate whorl is composed of 7 or more spherical chambers that increase slowly in size. The periphery is distinctly lobulate and the surface of the test is relatively smooth and evenly covered with small pustules. The axial periphery is imperforate but is pierced by occasional pores. The aperture is a high rounded umbilical-extraumbilical arch that is bordered by a thin, narrow lip.

Discussion: Liu and Olsson (1994) showed that a variety of morphotypes existed within an assemblage of H. monmouthensis. This species is regarded as ancestral to the normal perforate cancellate species of the earliest Danian which also have the general shape of these morphotypes. Thus, it is believed that these variations may have established trends that resulted in the separation of the Paleocene globigerinids and globorotaliids. Very primitive pore pits can be observed in the chamber walls of H. monmouthensis. In the earliest Danian this character was strongly developed, resulting in the cancellate wall texture which is the most important distinguishing feature of Cenozoic planktonic foraminifera. Two important cancellate groups of planktonic foraminifera are derived from H. monmouthensis, cancellate spinose and cancellate nonspinose. In the first instance, Eoglobigerina and Parasubbotina represent the first of the cancellate spinose genera and, in the second, Praemurica represents the first of the cancellate nonspinose genera. A discussion of these transitions is given in Liu and Olsson (1994).

Stable isotopes: Hedbergella monmouthensis has among the most positive 18O of any Cretaceous trochospiral planktic foraminifer (Boersma et al. 1979). The 18O of H. monmouthensis is similar to Abathamphalus mayaroensis and distinctly more positive than Rugoglobigerina rotundata, R. rugosa, Globotruncanella petaloidea, and Globotruncana arca (Berggren and Norris, in press). The 13C of H. monmouthensis is lighter than coexisitng Rugoglobigerina.

Stratigraphic range: Upper Maastrichtian to lower Zone P0.

Global distribution: Worldwide in high and lower latitudes.

Origin of species: Originates in the upper Maastrichtian from H. holmdelensis.

Repository: Holotype (USNM 626471, lost), topotype (USNM 640920), and neotype (USNM 488562) deposited in the Cushman Collection, U.S. National Museum. Examined by BTH and RKO.

Type species: Globanomalina ovalis Haque, 1956

Diagnostic characters: Test very low trochospiral with 12-15 chambers, and with 5-6 in the ultimate whorl. The chambers vary from globular to ovoid to low conical in shape. The aperture is interiomarginal, umbilical to extraumbilical, a moderately high arch which is bordered by a narrow lip that may broaden slightly towards the umbilicus. The aperture may extend slightly onto the spiral side. The umbilicus is small and generally deep. The wall is normal perforate and smooth but may be covered with pustule growth in some species, particularly in the umbilical area. The peripheral margin may be perforate, display an imperforate band, or be keeled.

Discussion: Banner's 1989 study of Globanomalina clarified its taxonomic status, clearly distinguishing the genus from the planispiral Pseudohastigerina which Loeblich and Tappan (1988) regarded as a junior synonym of this genus. Banner emended the genus to clearly separate it from Pseudohastigerina and discussed its evolutionary relationship with this genus. He, however, regarded Globanomalina as having a microperforate wall with low, bluntly margined perforation pits. His observations appear to be based on poorly preserved specimens where the original wall has been corroded and recrystallized. All of the species of Globanomalina included in this atlas are normal perforate. Banner's observation of perforation pits is, however, correct but as he pointed out they are not to be confused with a cancellate wall. Globanomalina is here emended to include the smooth-walled species of the Paleocene which are members of two distinct lineages, one with an imperforate margin which leads to the carinate species G. pseudomenardii and one with a perforate margin which leads to the planispiral genus Pseudohastigerina.

Globorotalia (Turborotalia) archeocompressa Blow, 1979, p. 1049, holotype: pl. 58, figs. 8-9 (Zone P , DSDP Hole 47.2/11/5: 148-150 cm); paratypes: pl. 56, figs. 1-2 (Zone P , DSDP Hole 47.2/11/6: 148-150 cm), pl. 58, figs. 2, 6,7 (Zone P , DSDP Hole 47.2/11/5: 148-150 cm), pl. 64, figs. 5,9, (Zone P , DSDP Hole 47.2/11/3: 148-150 cm), pl. 68, figs. 5,6 (Zone P1, DSDP Hole 47.2/11/3: 0-5 cm), Shatsky Rise, NW Pacific Ocean.

Original description: The very small test consists of about 11-12 chambers coiled in a low, lax trochospire with 5 1/2 chambers visible in the final convolution of the test. In dorsal aspect, the chambers are almost hemispherical with inflated dorsal surfaces and depressed, sensibly radially disposed, intercameral sutures; the chambers are almost equidimensional with the tangential length only very slightly greater than the radial breadth. The inflation of the dorsal surfaces of the last convolution of chambers is such that the initial spire of chambers is depressed below the level of the later chamber surfaces. In equatorial profile, the test is moderately lobulate but is almost circular in overall outline. In axial-apertural profile, the test appears biconcave because of the depression of the initial spire below the level of the dorsal surfaces of the later chambers. The test is also seen in the axial-apertural view to have a rounded peripheral marginal outline and the later chambers, at least, appear subglobular with little or no differentiation in the degree of inflation for the ventral and dorsal surfaces of the chambers. In ventral aspect, the chambers appear subglobular but with quite deeply depressed, but not sharply incised, intercameral sutures which are radially disposed. The umbilicus is open but is shallow with no clearly marked umbilical shoulders to the surrounding convolution of chambers. The aperture is interiomarginal and extends from the shallow umbilicus to very nearly to the peripheral margin. The aperture is bordered by a well-marked flange-like lip and is a low, only slightly arched, opening. The wall is very finely perforate and the mural-pores do not open into distinct pore-pits, neither are they densely arranged. In optical appearance, the finely and sparsely perforate wall combined with the absence of pore-pits and inter-pore ridges gives the test a smooth, almost polished appearance which is characteristic of the G. (T.) compressa (sl) lineage. Maximum diameter of holotype) 0.139 mm as measured by electron-beam sensor.

Diagnostic characters: This is a very small-sized species, being less than 130 um in greatest diameter. It has a flattened compressed test with the spiral side being nearly plane. The axial periphery is rounded with slightly conical chambers which are nearly flat on the spiral side and project at a high angle towards the umbilicus. Five to six, compressed oval-shaped chambers that increase very gradually in size make up the ultimate whorl. The umbilicus is shallow and broad, open to the previous chambers, and contains sparse pustules on the umbilical shoulders of the surrounding chambers. The umbilical-extraumbilical aperture is a broad, low arch which is bordered throughout its extent by a narrow lip. The wall, which ranges from 4 to 7 um in thickness, is smooth and perforate. The pores which average 1 um at the narrowest point are mostly confined to the chamber surfaces away from the periphery which is largely imperforate.

Discussion: This is the first species of the genus Globanomalina to appear in the earliest Danian. In morphology it is very similar to its ancestral species Hedbergella holmdelensis Olsson, 1964. It differs, however, in that fine pustules are confined to the umbilical and inner spiral areas of the test rather than being sparsely distributed over the entire walls of the test as in H. holmdelensis. In early species of Globanomalina the chamber walls are smoother and generally free of numerous pustules. Pustules are more numerous on the chamber walls of G. pseudomenardii but these are enhanced by gametogenesis. The possibility that the "pustules" in archeocompressa are of similar origin needs further investigation. The chambers are somewhat more inflated and the equatorial peripheral margin is more lobulate in H. holmdelensis. Another difference lies in the ontogeny of each species. Six to 7 chambers make up the penultimate whorl in holmdelensis, a chamber arrangement typical of ancestral species of Hedbergella in the Cretaceous. In archeocompressa the penultimate whorl is reduced to 5 chambers which is typical of Globanomalina. The imperforate peripheral margin is a primary derived taxonomic character from holmdelensis and is present throughout the archeocompressa lineage.

Stable isotopes: No data available.

Stratigraphic range: Upper Zone P0 - P1b. Due to the very small size and rare occurrence of this species the upper range of this species is not yet reliably established.

Global distribution: Western equatorial Pacific, Gulf coast, South Atlantic Oceans (DSDP Site 356).

Origin of species: This species evolved from Hedbergella holmdelensis in late Biochron P0.

Repository: British Museum of Natural History, holotype: British Petroleum Catalog No. 37/38, paratypes: BP Cat. No. 37/7,21,39,49,58,118; 40/12,44,55.

Globorotalia australiformis Jenkins, 1966, p. 112 , fig. 11, nos. 92-96 (lower Eocene, Waipawan Stage, Middle Waipawan River section, New Zealand). - Ludbrook and Lindsay, 1969, p. 367, pl. 1, figs. 4-5 (middle Eocene, South Australia)

Globorotalia (Planorotalites) australiformis Jenkins, in Hornibrook, Brazier and Strong, 1989, p. 128, fig. 24:15a-c (holotype refigured).

Planorotalites australiformis (Jenkins). -Tjalsma, 1977, p. 495, pl. 2, figs 10-12 to (lower Eocene, DSDP Site 329/32/1: 141-143 cm), fig. 13, (upper Paleocene, DSDP Site 329/33/1: 125-127 cm), Maurice Ewing Bank, South Atlantic Ocean. -Jenkins, 1985, p. 279, fig. 6.8 (holotype refigured). -Huber, 1991b, p. 440, pl. 6, figs. 17,18 (uppermost Paleocene, Zone AP5, ODP Site 738B/23X: 196.93 msbf, Kerguelen Plateau, Southern Indian Ocean).

Original description: Test free, sinistrally coiled, small, low trochospiral, biconvex, equatorial periphery quadrilobate; axial periphery rounded in the first 3 chambers of the final whorl, but angled with an incipient keel in the final chamber. Wall calcareous, finely perforate, surface smooth. Chambers compressed, 13 chambers in about 2 1/2 whorls, with 5 chambers in the first whorl and 4 in the last, increasing fairly rapidly in size; proloculus diameter approximately 0.01-0.02 mm. Sutures recurved on spiral side and nearly radial on the umbilical side; slightly depressed on both sides. Umbilicus small, open. Aperture a low arch interiomarginal, extraumbilical-umbilical. Maximum diameter: 0.26 mm.

Diagnostic characters: This small species is distinguished by having 4 low conical shaped chambers in the ultimate whorl and an acute axial periphery. The earlier chamber walls are covered by a fine, dense pustules which become more sparsely distributed on the penultimate and ultimate chambers. The axial periphery is imperforate for the most part and becomes thickened on the last one or two chambers.

Discussion: The original description of this species by light microscope indicated a smooth, finely perforate wall but SEM micrographs show a fine coating of pustule-like calcite covering the original wall. It is possible that this coating is of gametogenetic origin but this possibility needs more study. The species has been only reported in southern latitude sites and may be biogeographically restricted. Blow (1979) suggested that australiformis may be a cool-water endemic form of the tropical Globorotalia troelseni Loeblich and Tappan morphotype due to its general morphologic similarity. Globorotalia troelseni (=G. chapmani Parr, 1938) is a 5 chambered species and does not possess the dense pustulose wall surface of australiformis. It would appear that australiformis arose independently as an endemic species in the southern high latitudes. Blow also placed specimens identified as Globorotalia elongata Glaessner, 1937 by Loeblich and Tappan (1957a) from the Vincentown Formation of New Jersey and from the Velasco Shale of Mexico in synonymy with australiformis. These specimens belong to G. chapmani.

Stable isotopes: No data available.

Stratigraphic range: Lower part of Waipawan Stage to the Porangan Stage; upper Subbotina triloculinoides Zone to the P. primitiva Zone (upper Paleocene to lower Eocene).

Global distribution: Southern mid-high latitudes.

Origin of species: The origin of this species has not been traced. Jenkins (1966) notes that australiformis is closely related to many Paleocene smooth-walled species (of Globanomalina) and that it replaces G. pseudomenardii Bolli, 1957a in the lower part of the Waipawan Stage. It would appear more closely related to G. imitata in having 4-4 1/2 (mostly 4) chambers in the final whorl and in the dense pustulose wall surface observed in early chambers of this species.

Repository: N.Z. Geological Survey Register No. TF 1503: holotypes and two paratypes.

Globorotalia chapmani Parr, 1938, holotype: pl. 3, figs. 9a,b; topotype: pl. 3, fig. 8 [upper Paleocene (probably Zone P4), Kings Park Shale, Kings Park Bore 1, 1,755 ft., Perth Basin, W. Australia]. -McGowran, 1964, p. 85, text-figs. 1-9 (upper Paleocene, Carnarvon Basin, N.W. Australia). -Berggren, Olsson, and Reyment, 1967, p. 277, text-figs. 1, 3, nos. 1a-c, 4, nos. 1a-c, pl. 1, figs. 1-6 (Zone P4, Vincentown Fm., New Jersey; Boongerooda Greensand, N.W. Australia, text-fig. 1). -Pujol, 1983, p. 657, pl. 3, fig. 3 (upper Paleocene, DSDP Hole 516F/85/1: 40-42 cm, Rio Grande Rise, South Atlantic Ocean) .-Haig, Griffin and Ujetz, 1993, p. 275, pl. 1, figs. 1-3 (holotype ESEM), 4-6 (Parr topotype ESEM, from same level as holotype); pl. 2, figs. 1-23 (topotypes, from same level as holotype).

Anomalina luxorensis Nakkady, 1951, p. 691, pl. 90, figs.39-41 (upper Paleocene, Egypt).

Globanomalina ovalis var. lakiensis Haque, 1956, p. 149, pl. 14, figs. 2a-c (upper Paleocene, Pakistan).

Globorotalia membranacea (Ehrenberg). -Subbotina, 1953, (partim), p. 205, pl. 16, figs. 12a-c (not pl. 16, figs. 7a-11b, 13) (Upper Paleocene, Crimea, Tarkhankut peninsula).

Globorotalia elongata Bolli, 1957a (non Globorotalia pseudoscitula var. elongata Glaessner, 1937), p. 77, pl. 20, figs. 11-13 (Zone P4, Lizard Springs Fm., Trinidad).

Globorotalia troelseni Loeblich and Tappan, 1957a, p. 196, pl. 60, figs. 4a-c (Zone P4, Nanafalia Fm., Alabama), pl. 63, figs. 5a-c (Zone P4, Velasco Shale, Mexico).

Globorotalia (Globorotalia) ehrenbergi Hillebrandt (non Bolli, 1957a), 1962, p. 126, pl. 12, figs. 3a-c (Zone D, Paleocene, Reichenhall-Salzburg Basin, Austro-German border).

Planorotalites chapmani (Parr). -Nederbragt and Van Hinte, 1987, (partim),p. 586, pl. 2, figs. 3-10 (not figs. 1,2, 11-16), pl. 3, figs. 4-6 (not figs. 1-3, 10-15) (Zone P4, DSDP Site 605/46/4: 85-89 cm, fig. 3, 605/47/3: 60-62 cm), New Jersey margin) .-Huber, 1991b, p. 440, pl. 6, figs. 19, 20 (upper Paleocene, ODP Hole 738C/9R: 265.01 msbf, Kerguelen Plateau, Southern Indian Ocean).

Original description: Test biconvex, oval, the dorsal surface more convex than the ventral, which is umbilicate; periphery lobulated, peripheral margin rounded; chambers comparatively few, not more than five in the last-formed whorl, each much larger than its predecessor; sutures depressed, not limbate, gently recurved on both sides of the test; wall smooth and punctate, with a silvery lustre; aperture an elongate slit with a slight lip, opening at the base of the last-formed chamber into the umbilical depression. Length up to 0.65 mm. This species belongs to the group of G. hirsuta (díOrbigny) and is perhaps nearest to G. hirsuta, which typically has only four chambers to a whorl and with the sutures on the ventral side radial.

Diagnostic characters: This species is identified by its compressed test, pinched periphery with a thickened imperforate band, and the rapidly enlarging chambers. The number of chambers in the ultimate whorl is typically 5 but can range up to 6. The test walls are smooth with occasional small pustule buildups in the umbilical area and on the inner spiral area.

Discussion: Haig et al.'s illustrations of the holotype, paratypes, and topotypes by the SEM has clarified the morphologic characters of this species. It is a common species in upper Paleocene assemblages.

Stable isotopes: Globanomalina chapmani has 13O and 13C similar to Parasubbotina varianta, S. triloculinoides and G. pseudomenardii. The species has distinctly more positive 18O and more negativeº 13C than Morozovella, Acarinina, and Igorina.

Stratigraphic range: Within Zone P3 to Zone P5.

Global distribution: Worldwide in mid-latitude and high latitude sections.

Origin of species: Globanomalina chapmani is a member of the smooth-walled imperforate periphery lineage and evolved from G. ehrenbergi (Bolli) in the lower part of Zone P4.

Repository: Museum collections of the Department of Geology, University of Western Australia, UWA18897.

Globigerina compressa Plummer, 1926, p. 135, pl. 8, figs. 11a-c [Zone P2, Wills Point Fm., Midway Group (upper Danian), Navarro County, Texas].

Globigerina compressa var. compressa Plumme. -Subbotina, 1953, p. 63, pl. 2, figs. 2a-6c (Danian, Zone of rotaliform globorotaliids, Elburganian horizon, N Caucasus)

Globorotalia compressa (Plummer. -Bolli, 1957a, p. 77, pl. 20, figs.21-23 (lower Paleocene, Lizard Springs Fm., Trinidad). -Bolli and Cita, 1960, p. 20, pl. 32, figs. 3a-c (Gt. trinidadensis-Gl. daubjergensis Zone, Paderno d'Adda section, N Italy). -Pujol, 1983, (partim), p. 656, pl.2, figs. 3,4 (not fig. 2) (lower Paleocene, DSDP Site 516F/89/2:119-121 cm, Rio Grande Rise, south Atlantic Ocean).

Globorotalia (Globorotalia) compressa (Plummer). -Hillebrandt, 1962, p. 125, pl.12, figs. 1a-c (Zone B, lower Paleocene, Reichenhall-Salzburg Basin, Austro-German border).

Globorotalia (Turborotalia) compressa compressa (Plummer). -Blow, 1979, p. 1062, pl. 75, figs. 10, 11 (Zone P1, Lindi area, Tanzania, E Africa), pl. 78, figs. 5-10 (Zone P2, DSDP Hole 20C/6/4: 72-74 cm), pl. 248, figs. 1-3 (topotype illustrations), pl. 254, figs. 1-3 (Danian, Denmark), pl. 257, figs. 5-7 (upper Danian, Sweden)

Planorotalites compressus (Plummer). -Huber, 1991c, p. 461, pl. 3, figs. 1, 2 (Zone AP2, ODP Hole 738C/16: 340.93 mbsf (southern Kerguelen Plateau, Southern Indian Ocean).

Globanomalina compressa (Plummer). -Berggren, 1992, p.563, pl. 1, figs. 14-16 (ODP Hole 747A/19H/CC, Kerguelen Plateau, Southern Indian Ocean).

Original description: Test small, rotaliform, closely coiled, somewhat compressed, equally bioconvex; peripheral margin bluntly angular, lobate; chambers increasing gradually, 5 in last-formed whorl, moderately inflated, overlapping on dorsal face; sutures distinctly depressed and strongly curved on the dorsal side; shell wall thin, smooth, finely punctate; aperture a single moderately arched slit protected by a definite flaring flap at base of septal face and extending into the small but distinct umbilical depression.

Diameter up to .4 mm.; average .3 mm.

Diagnostic characters: A small, 5 chambered smooth-walled test, with a moderately angular axial periphery, and with an imperforate peripheral margin that is moderately to strongly developed. Aperture a low, umbilical-extraumbilical arch, bordered along its entire extent by a narrow well defined lip.

Discussion: Plummer (1926) designated three co-types to represent her new species and illustrated one view of each (dorsal, edge and ventral). We have selected one of these co-types (Plummer, 1926, fig. 11c) as a neotype for this species. The axial view of the neotype is the view illustrated by Plummer to show the degree of compression of the test and the bluntly angular peripheral margin of the chambers of her new species. The chambers of the two other co-types are somewhat more inflated and the axial periphery of the test is somewhat less compressed than is the neotype. Blow (1979) restricted his identification of G. compressa to morphotypes with compressed ogival chambers in axial view in contrast to morphotypes with an inflated rounded axial periphery which he identified as cf. compressa or, if the chambers were more fully inflated, as G. planocompressa Shutskaya. Our studies separate compressa and planocompressa on the degree of compression/inflation of the chambers and the presence or absence of an imperforate peripheral margin. In the compressa-ehrenbergi-chapmani lineage an imperforate peripheral margin is a distinguishing characteristic and the degree of compression of chambers in the ultimate whorl may vary from slightly compressed to the compressed ogival shape. In contrast, in the planocompressa-imitata-ovalis lineage the chambers in the ultimate whorl are fully inflated and the axial periphery is perforate.

Stable isotopes: Globanomalina compressa has 18O and 13C similar to Parasubbotina varianta, S. triloculinoides and G. ehrenbergi. The species has distinctly more positive 18O and more negative 13C than Morozovella and Praemurica.

Stratigraphic range: Zone P1c to P3.

Global distribution: Worldwide in low to high latitudes.

Origin of species: This species shares morphologic similarities with G. archeocompressa in the compressed test and in the bluntly angular imperforate peripheral margin. The upper range of archeocompressa has not been firmly established and, consequently, its linkage with compressa is, at present, uncertain. We link the two species on the basis of their shared morphologic similarities.

Repository: The three cotypes, Walker Museum Collection 33078, Station 23 are designated as follows: neotype, PE 55091, and paratype, PE 33078, Chicago Field Museum; paratype (USNM 488563), U.S. National Museum. Examined by BTH and RKO.

Globorotalia membranacea Cushman and Bermudez, 1949 (non Planulina membranacea Ehrenberg, 1854), p. 34, pl. 6, figs. 16-18 (Paleocene of Cuba).

Globorotalia ehrenbergi Bolli, 1957a, p. 77, pl. 20, figs. 18-20 (upper Paleocene, Lizard Springs Fm., Trinidad).

Globorotalia haunsbergensis Gohrbandt, 1963, p. 53, pl. 6, figs. 10-12 (Paleocene, N Salzburg, Austria).

Globorotalia (Turborotalia) haunsbergensis Gohrbandt. -Blow, 1979, p. 1075, pl. 88, figs. 6,8 (Zone P3, DSDP 6/47.2/10/1:72-74 cm, Shatsky Rise, NW Pacific Ocean), fig. 9 (Zone P3, Lindi area, Tanzania, E Africa).

Original description: Shape of test low trochospiral, compressed; equatorial periphery strongly lobate; axial periphery acute, last chamber often with a faint keel. Wall calcareous, perforate, surface smooth. Chambers compressed; about 12-15, arranged in 2-3 whorls, the 5 chambers of the last whorl increasing fairly rapidly in size. Sutures on spiral side slightly curved, distinctly depressed; on umbilical side radial, depressed. Umbilicus shallow, open. Aperture a low arch, with a lip; interiomarginal, extraumbilical-umbilical. Coiling random. Largest diameter of holotype 0.28 mm.

Diagnostic characters: Compressed, smooth-walled test, with chambers moderately increasing in size, often with the ultimate chamber smaller than the penultimate one, pinched periphery that has a thickened imperforate margin (faint keel of Bolli), 5-5 1/2 chambers in the ultimate whorl.

Discussion: Bolli (1957a) erected this species for Paleocene morphotypes that had been referred to Ehrenberg's species. The basis for distinguishing this species from G. compressa is the pinched periphery and the thickened imperforate margin which is more strongly developed than in compressa. Although Blow (1979) separated ehrenbergi from G. haunsbergensis we include morphotypes with a more sharply acute periphery within the range of variation of ehrenbergi. Blow drew attention to the more lax coiling of haunsbergensis morphotypes which results in a wide, shallow umbilicus. The trend towards a more strongly developed imperforate margin was recognized by Bolli and Blow as a trend towards a true keel which resulted in the evolution of G. pseudomenardii. Globanomalina chapmani is also allied to ehrenbergi in having a pinched periphery and a thickened imperforate margin. As in pseudomenardii the rate of chamber size increase is greater than in ehrenbergi.

Stable isotopes: Globanomalina ehrenbergi has 18O and 13C similar to Parasubbotina varianta, S. triloculinoides and G. compressa. The species has distinctly more positive 18O and more negative 13C than Morozovella and Igorina.

Stratigraphic range: Zone P2 to Zone P4.

Global distribution: Apparently a worldwide distribution in low to mid-latitude sections.

Origin of species: As pointed out by Bolli and Blow this species is transitional to G. pseudomenardii. It evolved from G. compressa by developing a more highly developed imperforate margin and a more sharply angular axial margin.

Repository: Holotype (USNM P5060) deposited in the Cushman Collection, U.S. National Museum. Examined by RKO.

Globanomalina imitata (Subbotina), 1953

Original description: Shell small, rotaliform, strongly convoluted with a broadly oval outline. Whorls 2 1/2-3. The first 2 whorls are considerably smaller than the last. Their diameter comprises on average only one third the diameter of the whole shell. Dorsal side flattened, ventral side convex, subconical. In the centre of the central side there is a very small, hardly discernible umbilicus. The umbilical ends of the chambers are disposed above the rest of the surface of the ventral side, thus forming the apex of a short cone. The peripheral margin is rounded and scalloped. The chambers on the dorsal side are oval and drawn out in the direction of the spiral whereas on the ventral side they are broadly triangular. There are 4 chambers to the last whorl and these show a rapid increase in dimensions. The sutures are recessed, almost straight or slightly curved, on both the dorsal and the ventral sides. Attention should be paid to the shortness of the sutures on the dorsal side as compared with the rather long sutures on the ventral side. The orifice is represented by a straight slit covered by well-defined slender lips. It has the form of an even rectangular cavity extending along the whole length of the marginal suture. The wall is thin, smooth, and in well-preserved specimens lustrous; it is perforated by many small pores. Dimensions: diameter 0.15-0.25 mm, thickness 0.08-0.11 mm.

Diagnostic characters: A small smooth-walled test with 4 to 4 1/2, occasionally 5, inflated chambers in the ultimate whorl. The chambers are ovoid in shape with the long axis directed towards the umbilicus. In spiral and umbilical view the chambers are ovoid in shape with the long axis parallel to the coiling spire. The test walls are perforate throughout and the aperture is a high umbilical-extraumbilical arch that is bordered by a thin continuous lip. The wall is smooth in the adult chambers, but is covered with fine scattered to dense pustules in the early ontogenetic stages which possess anguloconical chambers.

Discussion: This is a distinctive species which is often overlooked because of its small size. It belongs to the perforate walled, inflated chamber lineage of planocompressa-imitata-ovalis. It also appears to have a phylogenetic link to G. australiformis (see discussion under this species). The specimens illustrated by Blow (1979) as imitata are distinctly cancellate walled and do not belong to this species but probably to Praemurica.

Stable isotopes: No data available.

Stratigraphic range: Zone P1c to P4.

Global distribution: Known from the northern Caucasus, southeast India, and the Atlantic and Gulf Coastal Plains.

Origin of species: Probably evolved from Globanomalina planocompressa in Zone P1c.

Repository: St. Petersburg VNIGRI (378\112), holotype No. 4073, paratypes No. 4074, 4075. Examined by FR.

Globanomalina ovalis Haque, 1956, p. 147, pl. 14, figs. 3a-c (upper Paleocene, Bed B2, Laki Fm., Pakistan)-Banner, 1989, emended, p. 177, pl. 1, figs. 1a-c (original drawings of holotype), figs. 2a-c (original drawings of holotype of Globanomalina simplex Haque, 1956), 3a-c (SEM of paratype of G. ovalis), 4a-c (original drawings of holotype of Globigerina pseudoiota Hornibrook, 1958); pl. 2, figs. 1a-c (paratype of G. ovalis), figs. 2a-e (upper Paleocene, Dannevirke Series, New Zealand); pl. 3, figs. 1a-2d (upper Paleocene, Dannevirke Series, New Zealand)

Globorotalia (Turborotalia) sp. ex interc G. (T.) chapmani Parr and Pseudohastigerina wilcoxensis (Cushman and Ponton), Blow, 1979, pl. 111, fig. 5 (Moogli Mudstones, Papua, given as Zone P7 which is correlative with Zone P6a as used in this atlas).

Globigerina pseudoiota Hornibrook, 1958, p. 34, pl. 1, figs. 16-18 (upper Paleocene of New Zealand)

Original description: Test small, oval, longer than broad, biconvex, nearly bilaterally symmetrical. Periphery sub-acute, broadly rounded, lobate. Dorsal side flat or slightly convex, showing all the chambers. Ventral side completely involute, umbilicate. Chambers distinct, inflated, globose, somewhat overlapping, about 6 in the adult whorl, rather rapidly increasing in size as added; last 2 chambers occupy more that half of the test. Sutures distinct, depressed, somewhat curved on the dorsal side, radial on the ventral. Surface smooth, shiny. Wall calcareous, very finally perforate, built of radially arranged crystals of calcite. Aperture an elongate slit at the base of the last chamber of the outer whorl extending from the periphery to the umbilicus with a narrow lip. Length of figured specimen 0.26 mm; breadth 0.21 mm; thickness 0.14mm.

Diagnostic characters: emended description- Globanomalina as here defined, with six chambers per whorl initially, sometimes retaining this number in the last-formed whorl but sometimes reducing it to five; the rate of chamber volume increase with chamber addition is approximately the same in each form, so that (a) when the chamber number is reduced (e.g., to five in the last whorl) the spiral side of the test remains completely evolute and flat or slightly convex in shape, or (b) when the final chamber number is not reduced (e.g., is retained at about six in the last whorl) the final chambers overlap slightly onto the spiral side, so that that side becomes incompletely evolute and slightly concave in shape; wall is microperforate, with perforation canaliculi about 1 um in diameter, opening into broadly and bluntly rounded perforation pits about 10 um in diameter (the perforation pits are deeper on the earlier chambers which have had their walls thickened by test growth); maximum diameter achieved is about 0.34 mm.

Discussion: Banner's (1989) study has clarified the taxonomic positon of the genus Globanomalina and the type species ovalis. Banner correctly points out that as the number of chambers increase from 5 to 6 in the ultimate whorl the test becomes asymmetrically planispirally arranged. In addition, the aperture which is a fairly high arch, bordered by a continuous, thin lip, migrates slightly onto the spiral side. The test walls are perforate throughout and, although Banner describes the wall as microperforate, we include ovalis in the normal perforate category which contrasts with microperforate taxa (test walls with pore diameters 0.3 to 0.45 um in diameter) of the Guembelitridae.

Stable isotopes: No data available.

Stratigraphic range: Upper Zone P4 to Zone P6.

Global distribution: Northern and southern mid-latitudes.

Origin of species: This species is a member of the inflated chamber, perforate walled lineage in Globanomalina. Globanomalina ovalis evolved from G. imitata in the upper part of Zone P4 by an increase in the number of chambers in the ultimate whorl from 5 to 6, a greater degree of inflation of the chambers, and the trend towards planispirality. Although Berggren et al., 1967 and Banner, 1989 derive ovalis from G. chapmani, there is a distinct difference in wall texture between the two species that would appear to preclude a phylogenetic linkage. In the Pondicherry section of southeast India ovalis and imitata overlap morphologically.

Repository: Geological Survey of Pakistan at Quetta, Pakistan. Paratypes at British Museum (Natural History), specimens P.42400.

Globorotalia planocompressa planocompressa Shutskaya, 1965, p. 179, pl. 1, figs. 6a-c (lower Paleocene, N Caucasus).

Globorotalia compressa (Plummer). -Loeblich and Tappan, 1957a (partim), p. 188, pl. 40, figs. 5a-c (type Danian, Ostratorp), (paratypic by synonymy); pl. 41, figs. 5a-c (Zone P1b, McBryde Fm., Alabama); pl. 42, figs. 5a-c (Zone P1b, Brightseat Fm., Maryland)

Globorotalia (Turborotalia) compressa planocompressa Shutskaya. -Blow, 1979, p. 1067, pl. 68, figs. 4, 8-10 (Zone P1a, DSDP Hole 47.2/11/3: 0-5cm), pl. 71, figs. 8-10 (Zone P1, DSDP Hole 47.2/1/1:0-1cm, Shatsky Rise, NW Pacific Ocean).

Globanomalina planocompressa Shutskaya. -Olsson, Hemleben, Berggren, and Liu, 1992, p. 207, pl. 7, figs. 6-8 (Zone P1a, Clayton Basal Sands, Millers Ferry, Alabama).

Original description: Test with a subpolygonal lateral outline, strongly compressed along the growth axis. Spire consists of three whorls. Early whorls lie on one level with the surface of the last whorl or rise up slightly above it. In each whorl there are five to six inflated subspherical chambers, rapidly and uniformly increasing in size and arranged freely. Peripherally on the chambers a small papillate projection, directed toward the subsequent chamber, is frequently observed. Last chamber asymmetrical, flattened at the apertural face and slightly shifted toward the umbilical surface. Equatorial outer margin scalloped. Axial outer margin weakly asymmetric, broadly rounded. Sutures deep, straight. Umbilicus narrow. Aperture a semicircular slit located between the umbilicus and the outer margin. Sometimes it is surrounded by a very narrow lip, the width of which is uniform over