NUMBE 8R5 77 47.2/10/5 :70-72 cm] ,pi .77 :figs .2-5 [Zone P2 ,DSDP Hole 20C/6/4: reported in the literature from predominantly low latitude 72-7 4cm B; raz iBl asin S, outh Atlanti cOcean] p, i8. 2 f:igs 1. - 3[Zone P2, recollectio no Gf loborotali auncinat atyp elocalit yT,rinidad ]p, i8.4 f:ig 2., (sub)tropical localities. Shutskaya (1970a, 1970b) recorded it pi .85 :fig .9 [Zone P3 ,DSDP Hole 47.2/10/2 :80-82 cm ;Shatsky Rise, from several localities in the northern Caucasus. This species northweste rPnaci fOiccea n[N], Motorozov 1a9,61.] does not appear to occur in high southern latitudes (Stott and Morozovel luancina t(aBolli).—Snyd earn Wdater 1s9, 85:448,44 p9 1,i .0fi:gs. Kennett, 1990; Huber, 1991b), and we have not found it in our 1 ,2 [Globorotalia uncinata Zone ,DSDP Site 550/37/5 :59-61 cm; examination of material from the southern part of the Indian Porcupi nAebys sPalla inno,rtheaste Arntlan Otic ean], Ocean (Figure 30). Original Description. —“Shape of test low trochospiral, ORIGIN OF Species .—This species evolved from Praemurica spiral side almost flat or slightly convex, umbilical side inconstans at the base of Zone P2 by extension of the distinctly convex; equatoria lperiphery distinctly lobate; axial umbilically conical chambers into most of the final whorl, and periphery rounded to subangular .Wal lcalcareous ,perforate, by the formation of blunt pustules around the umbilicus and on surface finely spinose .Chambers subangular ,inflated ,laterally the initia lchambers of the fina lwhorl. compressed; 12-15, arranged in about 2'/2 whorls, the 5-6 Repository. —Holotype (USNM P5048) deposited in the chambers of last whor lincreasing moderately in size. Sutures Cushman Collection, National Museum of Natural History. on spiral side strongly curved, depressed; on umbilical side Examined by WAB and RDN. radial ,depressed .Umbilicus fairly narrow ,deep ,open .Aper¬ ture a low arch; interiomarginal, extraumbilical-umbilical. Family Guembelitriidae Montanaro Gallitelli, 1957 Coiling random. Largest diameter of holotype 0.35 mm.” (Boll i1, 957a:74.) (by S. D’Hondt, C. Liu, and R.K. Olsson) Diagnostic Characters. —Nonspinose, weakly cancel- Guembelitrii nMaoentan aGraollit e1l9li5, 7: 1[s3u6bfamily], late ,elongate-oval ,plano-convex to moderately high spired, Guembelitriid Eale-Nagg a1r9,71:4 [3n1om ternanslat u semuxbfamily]. moderately lobulate test with 5-8 chambers in last whorl; Original Description. —“Test triserial; chambers globu¬ chambers occasionally so loosely coiled as to form secondary lar; aperture basal, arched, simple.” (Montanaro Gallitelli, spira lapertures between them ;sutures on umbilica lside radial, 1957:136.) depressed ,on spira lside incised and strongly recurved yielding Diagnostic Characters. —Original description of fo- typicall ytrapezoidal-shaped chambers ;axia pl eripher ysuban¬ raminifera ltests assignable to subfamily Guembelitriinae does gular ,noncarinate bu twith rugose muricae often situated along not apply to all members of family Guembelitriidae. Tests of peripheral margin of early chambers of last whorl; umbilicus guembelitriid foraminifera either triseria l (Guembelitria spp.), typically narrow, deep, bordered by weakly developed circu- trochospiral ( Parvularugoglobigerina spp., Globoconusa mumbilica lshoulder formed by raised periumbilica lchamber spp.), or nearly triserial in initial whorl and approximately extensions; aperture a narrow interiomarginal, umbilical- biserial in later whorls ( Woodringina spp.). Chambers usually extraumbilica larch extending to periphera lmargin. globular or subglobular ,increasing gradually in size .Aperture Discussion. —Blow (1979) argued the case for including usually a loop-shaped arch ,often slightly infolded on one side, uncinata Bolli, 1957a, as a junior synonym of praecursoria marked by a fine lip .Surface texture microperforate ,smooth to Morozova, 1957. Our examination of the holotypes of both of pustulous ;when present ,pustules or smal lmounds generally these forms leads us to reject this interpretation. Rather, we perforated by one or more pores (“pore-mounds”) ( Guembe¬ interpret praecursoria as an advanced, atypically large end- litria ,Parvularugoglobigerina ,Woodringina) or peripherally member of inconstans that is characterized by rounded associated with pore s (Globoconusa ). chambers and an axia lperiphery ,and we reserve for uncinata DISCUSSION. —Montanaro Gallitelli (1957) emended the those forms exhibiting the typica languloconica l(but noncari¬ family Heterohelicidae Cushman, 1927a, by creating the nate) test with distinctly incised and recurved spiral inter- subfamily Guembelitriinae fo rthe triseria lgenera Guembelitria cameral sutures. Acarinina indolensis Morozova (1959) is a Cushman, 1933, and Guembelitriella Tappan, 1940. Loeblich small form of P. uncinata with five chambers in the final whorl and Tappan (1957a) assigned Woodringina Loeblich and but ,nevertheless ,exhibits the characteristic morphology o fthis Tappan ,1957 ,to the Guembelitriinae ,implicitly modifying the species. definition of the Guembelitriinae to include forms that are Stable Isotopes. — Praemurica uncinata has more positive triserial in the first whorl and biserial in later whorls. El-Naggar 8 13 C and more negative 5 18 0 than Subbotina and Globanomal- (1971) raised the Guembelitriiidae to family status, and Blow ina and has an isotopic signature similar to that of Morozovella (1979) explicitly broadened its definition to include mor- praeangulata (Shackleton et al., 1985; Berggren and Norris, photypes with biseria land trochospira lstages. 1997). There is a pronounced increase in 8 13 C with increased The assignment of biserial Woodringina species to the size in P. uncinata (Kelly et ah, 1996; Norris, 1996). Guembelitriidae rests on the hypothesis that Woodringina Stratigraphic Range. —Zone P2 to lower Zone P3. species were derived from Guembelitria cretacea Cushman, Global Distribution. —This taxon has been widely 1933. This phylogenetic hypothesis is a subset of the broader 78 SMITHSONIA CNONTRIBUTION TS POALEOBIOLOGY 180°W 135°W 90°W 45°W 45°E 90°E 351°E FIGUR 3E 0—. Paleobiogeograph mic a sphowin dgistributio onPfraemuric uancina t(aBol liZin)on Pe2. hypothesis that Chiloguembelina midwayensis (Cushman, Among the primary morphologic characters that unite the 1940), is lineally derived from Guembelitria cretacea Cush¬ Guembelitriidae is a close surface-textura lassociation o fpores man, 1933, via Woodringina claytonensis Loeblich and and pustules (Liu and Olsson ,1992; Olsson et al. ,1992) (Plates Tappan, 1957 (Olsson, 1970, 1982; Premoli Silva, 1977; Li and 63-68) .Where the pores directly perforate the surface mounds Radford, 1991; Olsson etal., 1992; Liu and Olsson, 1992). This (o rpustules) ,these perforated mounds are commonly referred broader hypothesis is supported by cladistic analysis ,based on to as “pore-mounds.” Interspecimen and intraspecimen vari¬ shared apertural, surface textural, and juvenile characters ation in surface texture from smooth to pore-mound is (D’Hondt ,1991) .Given the apparen tphylogenetic relationship exhibited by Guembelitria cretacea ,Woodringina claytonen¬ o fChiloguembelina midwayensis to Guembelitria cretacea ,the sis ,and Parvularugoglobigerina species (D’Hondt and Keller, Guembelitriidae constitutes a paraphyletic family because it 1991; Liu and Olsson, 1992; Olsson et al., 1992) (Plates does not include descendent species assigned to the Chi- 63-68). The degree of pore-mound expression within a loguembelinidae. The relationship of many post-Paleocene specimen appears ontogenetically variable ,with the ultimate microperforate taxa (i.e., Cassigerinella Pokorny, 1955, Cor- chambe roften exhibiting more weakl ydeveloped pore-mounds rosina Thalmann, 1956, and Gallitellia Loeblich and Tappan, than immediately preceding chambers (Liu and Olsson ,1992) 1986 )to the Guembelitriidae remains largely unexplored. (Plates 63-68) .Furthermore ,the pustules (pore-mounds) are Recen sttudie shav edocumente dmorphologi cintergradation often asymmetrically perforate (Plate 63: Figure 6, Plate 65: and shared surface-textura land apertura lcharacteristics be¬ Figure 6, Plate 67: Figure 14). tween Guembelitria cretacea and members o fthe trochospiral The phyletic relationship of Parvularugoglobigerina, Glo¬ genera Parvularugoglobigerina Hofker ,1978 ,and Globocon- boconusa W, oodringina a, nd Chiloguembelina to Guembelitria usa Khalilov, 1956 (D’Hondt and Keller, 1991; Liu and Olsson, cretacea indicates that both trochospira land biseria lchamber 1992; Olsson et al., 1992) (Plates 63-68). These demonstrate arrangements divergently evolved within the planktonic fo- the genera lvalidity of severa lrecent phylogenetic hypotheses raminifera (Olsson, 1982; Smit, 1982; D’Hondt, 1991; Liu and for the origin of the latter genera (Olsson, 1970, 1982; Premoli Olsson, 1992). Such relationships have clearly not been Silva, 1977; Smit, 1982; Olsson et al., 1992). On this basis, we accounted for by taxonomic schemes ,which separate seria land assign both Parvularugoglobigerina and Globoconusa to the trochospiral morphotypes at the superfamily level (i.e., Guembelitriidae. Loeblich and Tappan ,1988). NUMBE 8R5 79 Genus Guembelitria Cushman, 1933 Guembelitria azzouz iSalaj ,1986:49 ,pi .1 :figs .1-6 ,pi .2 :fig .1 [base of Chiloguembe lHitroi af1k9e7r,8:60. Dania nHEl,ar iTau, nisia].Guembelitri abesbes Siala j1,986:50 p, i1 .f:igs 7. - 9[bas eo Df anian E ,Hl aria, TYPE Species .— Guembelitria cretacea Cushman ,1933. Tunisia], Original Description. —“Test similar to Gumbelina, but Guembelitr itarifol i(aMorozova).—Kelle r1,989:31 9fi,g s3.- 33,- [4lowerDanian B, razo sRive rT, exas],—D’Hond at n dKelle r1,991:93 p, i3. f:ig 2. triserial ;wal lcalcareous ,finely perforate ;aperture large ,at the [Zon ePa D, SD PSit e577/12/5 1:15-11 7cm S;hatsk yRise n,orthwestern inner edge o fthe last-formed chamber.” (Cushman ,1933:37.) Pacif Oiccean []N, oGtlobiger iEn (oaglobigerin atr)ifol iMaorozov a1,961.] Diagnostic Characters. —-Test small and triserial. Aper¬ ture bordered by a distinct lip and often slightly asymmetric. Original Description. —“Test small, triserial; chambers Wall structure microperforate; surface texture of well- globular, nearly spherical; sutures much depressed; wall preserved specimens characterized by presence of pore- smooth, finely perforate; aperture large, semicircular or mounds. semi-elliptica lat the inner margin of the last-formed chamber. DISCUSSION —. Confusion remain sove rthe taxonomi cstatus Length of holotype 0.20 mm.; breadth 0.17 mm.” (Cushman, of Chiloguembelitria, which Loeblich and Tappan (1988) 1933:37.) retained as a valid genus and Kroon and Nederbragt (1990) Diagnostic Characters. —Test small, triserial, composed suggested is a junior synonym of Guembelitria .This confusion o fglobula rchambers with strongly depressed sutures .Aperture is exacerbated by the absence of a designated holotype for bordered by distinct lip ;aperture often slightly asymmetric due Chiloguembelitria or its type species, Chiloguembelitria to infolding of lip along one side of aperture (as in danica Hofker ,1978 .Nonetheless ,it appears that no morpho¬ Woodringina and Chiloguembelina) .Wal lstructure microper¬ logica lcharacters consistently distinguish Chiloguembelitria forate; surface texture often characterized by blunt pore- spp. from Guembelitria spp. The original description and mounds; i.e., mounds marked by one or more pores (Plate 63: illustration of C .danica Hofker ,1978 ,resembles Guembelitria Figur e6). cretacea Cushman ,1933 ,in its apertura lcharacteristics and its DISCUSSION .—As discussed previously ,Chiloguembelitria consistent triseriality (Plate 63). The absence of distinct danica Hofker, 1978, appears to be a high-spired form of chiloguembelitriid characters indicates that ,as suggested by Guembelitria cretacea Cushman ,1933 .Similarly ,the primary Kroon and Nederbragt (1990), Chiloguembelitria danica characters of Guembelitria irregularis Morozova, 1961 (Plate Hofker, 1978, is a junior synonym of Guembelitria cretacea 8: Figures 1-3), and Guembelitria azzouzi Salaj, 1986, are Cushma n19, 33. indistinguishable from those of Guembelitria cretacea. In contrast, Guembelitria dammula Voloshina, 1961 (Plate 12: Guembelitria cretacea Cushman ,1933 Figures 7-9), appears to differ from Guembelitria cretaceasensu stricto by its three-chamber series being aligned more Figure 31 ;Plate 8 :figures 1-3 ;Plate 13 :figure 3; strictly parallel to its axis of test coiling. Closer examination of Plate 63 :figure s1-12 the type population of G. dammula may be necessary to Guembelitri acretace aCushman 1,933:37 p, i4 .f:ig 1.2a, b[uppe Mr aas- determine whether it should be reduced to a junior synonym of trichtian N, avarr oFm T„exas],—Olsson 1,970:601 p, i9.1 f:ig s4 .5[,upper G .cretacea or retained as a separate taxon. Maastrichtia Rne, dba nFkm N.,e Jwersey],—Smi tahn Pdessagn 1o9, 73:15, Salaj (1986) designated a short-spired earliest Paleocene p i1. f:igs 1. - 8[uppe rMaastrichtian C, orsican aFm .T, exas],—Keller, 1989:31 9f,ig s1.- 11,- [2lowe Dr anian B,razo Rsive rT,exas],—D’Hondt, form of Guembelitria as G. besbesi. Other authors have 1991:172 ,pi .1 :figs .1 ,3 ,5 ,6 [Zone Pa ,DSDP Site 577/12/5 :94-96 cm; identified that short-spired form of Guembelitria as Guembe¬ Shatsk Ryis en,orthwester nPacif iOc cean ]fi,g s 2[4.l,owe Dranian B,razos litria (?) trifolia (Morozova) or Guembelitria trifolia (Mo¬ River ,Texas] ,pi .2 :fig .2 [Zone Pa ,DSDP Site 577/12/5 :94-96 cm; rozova) (Blow, 1979; Keller, 1989; D’Hondt, 1991). The latter Shatsk yRise n, orthwestern Pacifi cOcean] f,ig 3.[Zon ePa D, SDP Site designations are incompatible with our interpretation of 528/31/CC 1:4-1 c5m W; alv iRsidg eS,out Ahtlant iOc cean].—D’Hondt and Keller 1, 991:93 p, i3. :fig 1. [Zone Pa D, SDP Site 577/12/5 :115-117 Globigerina (. Eoglobigerina ) trifolia Morozova, 1961, as a cm S;hats kRyis ne,orthweste rPnaci fOiccean].—L iaun Odlsso n19, 92:341, variant of Globoconusa daubjergensis (Bronnimann, 1953) pi .1 :figs .1 ,2 [Zone Pa ,Pine Barren Mbr. ,Clayton Fm. ,Millers Ferry, (Plate 8: Figures 4-6). In any case, the short-spired Guembe¬ Alabama]. litria morphotype intergrades with G. cretacea sensu stricto Guembelitr iarregular Mis orozov a1,961:1 7p 1,fi i.: [g9D. ania nT,arkhankut, (Plate 63). The former differs from the latter only in having a Crimea].—D’Hond t1,991:172 p, i1 .f:ig 7[.Zon ePa D, SD PSit e577/12/5: 94-9 c6m S;hatsk Ryis en,orthweste rPnacif Oiccean]. shorter spire. Along with G. cretacea sensu stricto, the Chiloguembelitr idaanic Ha ofke r1,978:6 0p, 4 if.:ig 1 .[4holotype D: anian, short-spired morphotype i spresen tin uppermos tMaastrichtian DSD PHol e47.2 s,ampl edept hunknown S;hatsk Ryise n,orthwestern near-shore sequences (D’Hondt, 1991) and is abundant in Pacifi cOcean].—Loeblich and Tappan 1, 988:452 p, i .484 :figs .3-6 lowermost Paleocene planktonic foraminifera lassemblages [holotype ]f,igs 7 . 8[,Danian D, SD PHol e47.2 s,ampl edept hunknown; (D’Hondt and Keller ,1991). The stratigraphic and biogeogra¬ Shats Rkyi sneo, rthweste Prnac iOficcean], Guembelitr i(a? t)rifol i(aMorozova).—Blow 1,979:138 4p ,6i.1 f:ig [9.Zone phic association of these intergrading morphotypes suggests P aD,SD HPo l4e7.2/11/ 51:48-15 c0m S;hatsk Ryis en,orthwester nPacific that the short-spired form (= Guembelitria besbes iSalaj ,1986) Ocean []N, oGtlobigerin (aEoglobiger itn r)aifo lMiaorozov 1a9, 61.] is a morphologic variant of G. cretacea. 80 SMITHSONIA CNONTRIBUTION TS POALEOBIOLOGY 180°W 135°W 90°W 45°W 45°E 90°E 135°E FIGUR 3.E—1 Paleobiogeograph mica sphowin dgistributio oGnfuembelitr ciaretac eCaushma tinh Deaman. Stable Isotopes. —Biogeographic and oxygen isotopic TYPE Species. — Globoconusa conusa Khalilov, 1956 = data suggest that G .cretacea inhabited a near-surface plank¬ Globigerin adaubjergensi sBronmmann 1, 953. tonic niche (Boersma et al., 1979; Boersma, 1984a; D’Hondt Original Description. —“Test high conical, turret-like, and Zachos 1, 993). tapering toward the initial end. Dorsal side strongly convex, Stratigraphic Range. —Maastrichtian to Zone Plb. with a conical spire, its apex at the initial chamber. The height Global Distribution. —During the Maastrichtian, G. of the cone exceeds the diameter of its base or is nearly equa lto cretacea was generally limited to near-shore environments and it. On the dorsal side all whorls are visible, in general only rarely was present in the open ocean (Davids, 1966; represented by semispherical chambers; on ventral side Olsson, 1970; Smith and Pessagno, 1973). Immediately subspherical chambers are observed only in the last whorl.” following the Cretaceous/Tertiary boundary event ,G .cretacea (Khalilov ,1956:249 ;translated from Russian). radiated into the open ocean ,where it was abundant at low and Diagnostic Characters. —Test tiny, low to high trochos- middle latitudes (Olsson, 1970; Smit and van Kempen, 1987; pire of 2—2 '/2 whorls; each whorl comprised of 3-4 inflated, D’Hondt and Keller, 1991; Liu and Olsson, 1992). In Zone P0 subglobular chambers .Wal lcalcareous ,microperforate ,with and lower Zone Pa it was a cosmopolitan planktonic distinctly hispid surface ;pustule soften peripherally associated foraminifera ltaxon .Biogeographic and oxygen isotopic data with a small pore (and vice-versa). Aperture a very small, low suggest that G. cretacea inhabited a near-surface planktonic arch ,with umbilica lposition .Test sometimes marked by one or niche (Boersma et al., 1979; Boersma, 1984a; D’Hondt and more smal lsupplementary apertures on spira lside. Zachos ,1993) (Figure 31). DISCUSSION. — Globoconusa conusa Khalilov, 1956, is Origin of Species.—U nknown. widely considered to be a junior synonym of Globigerina Repository. —Holotype (USNM CC19022) deposited in daubjergensis Bronnimann ,1953 (Loeblich and Tappan ,1964, the Cushman Collection ,Nationa lMuseum of Natura lHistory. 1988; Khalilov, 1967; Morozova et al., 1967, Olsson, 1970; Examined by SD, CL, and RKO. Premoli Silva, 1977; Hofker, 1978; Smit, 1982; Toumarkine and Luterbacher ,1985 ;Brinkhuis and Zachariasse ,1988 ;Stott Genus Globoconusa Khalilov, 1956 and Kennett, 1990; D’Hondt and Keller, 1991; Huber, 1991b;Li and Radford, 1991; Liu and Olsson, 1992; Olsson et al., Globast Biclao 1w97, 9:1231. 1992). A different opinion was held by Blow (1979), who noted NUMBE 8R5 81 that Khalilov’s (1956:249) type description of Globoconusa Diagnostic Characters. —Test characterized by conusa differs from the holotype of G. daubjergensis in some microperforate wall structure, with outer surface covered by respects (i.e., the test of G. conusa was originally described as abundant small pustules. Wide variation in spire height; some “strongly thickened” and “covered with small ,dense pits”) .On specimens marked by relatively low trochospire ,others display this basis, Blow (1979) created the generic designation relatively high spire (similar to short-spired Guembelitria Globastica for Globigerina daubjergensis and related forms. cretacea) .Occasionally exhibit ssmal lsecondary (intercameral) apertures; both primary and secondary apertures sometimes covered b ybullae. Globoconusa daubjergensis (Bronnimann ,1953) DISCUSSION. —The morphology of G. daubjergensis has Figure 32 ;Plate 8 :figures 4-6 ;Plate 15 :figures 13 ,14; been described in detai lby severa lauthors (Bronnimann ,1953; Plate 64 :figure s1-12 Loeblich and Tappan, 1957a; Troelsen, 1957; Blow, 1979). Globigerin daaubjergens iBsronniman n1,953:34 0te, xt-fi g[1D. anian, Troelsen (1957:120) recognized two forms as end-member Daub jQeruga Drreyn, mark], morphologic variants of Globigerina daubjergensis Bronni¬ Globoconus caonus Kahalilo v1,956:24 9p 5,fii.: g2.a -[cDania n,ortheastern mann, noting that the two variants “grade imperceptibly into Azerbaidzhan],—Blow 1,979:1386 p, i2.57 f:ig 1. 0[Danian D, SD PHole each other.” One of these contains four chambers in each whorl, 20C/6/4 7: 2-7 4cm B; raz iBl asin S, out hAtlanti cOcean ]f,ig 1. 1[Thanet whereas the other is a three-chambered form ( Eoglobigerina Sand Rse,culv eKre, nEtn,gland], Globigerinoid deasubjergen s(Bisronnimann).—Loebl iac nhTadppan, trifolia Morozova, 1961, Globigerina tripartita Morozova et 1957a 1: 84 p, i4. 0 f:ig l.a- c[uppe rDanian S, weden ]p, i4. 0 f:ig 8. a- c[Zone al., 1967, and Globastica sp. Type 1 of Blow, 1979). Blow PI P, ine Barren Mbr .C, layton Fm .A, labama] p, i4. 1 f:ig 9. a- c[Zone Pic, (1979:1244) suggested that the three-chambered form “is one McBryde Mbr. ,Clayton Fm. ,Alabama] ,pi .42 :figs .6a-7 c[Zone Pic, of three later differentiates from the basic Globastica daubjer- Brightsea Ftm .M, aryland ]p, i4. 3 f:ig l.a- c[Zon eP IK, incai dFm .T, exas], ge/ww-morphotype.” Close examination o flowermost Paleo- p i4. 4 f:igs 7. 8, a- c[Zon eP2 W, ill sPoin tFm .T, exas],—Olsson 1, 960:43, p i8. f:igs 4. - 6[Zon eP Ib, asa Hl omerstow nFm .N, ew Jersey]. cene sequences ,however ,suggests that the first appearance of Globigerin kaozlowsk Biirotze nan dPozarysk a1,961:16 2p , 1if.:ig s1.-1 4p,i. the three-chambered morphotype preceded that of the four- 2 f:igs 1. -17 p, i3 .f:igs l.a-2 c[Danian P, amietowo P, oland]. chambered morphotype (D’Hond tand Keller ,1991). Globigerin (aEoglobigerin t)arifol iMa orozov a1,961:1 2p , 1if.:ig [1D. anian, Some specimens assigned to the three-chambered mor¬ Tarkh aCnrikmute,a], photype appea rmorphologicall yintermediate between Guem¬ Globoconus atripartit aMorozov ae at l .1,967:193 p, i5 .f:igs 4. - 6[Danian, Kopet- CDraimghe,a]. belitria cretacea and Globoconusa daubjergensis (Plate 63: Globoconus daaubjergens igsigante Baang 1,969:6 5p , 4if.:ig s—1. 3 b[type Figures 7-9). The existence of such intermediate forms in D aDneiannm, ark], lowermost Paleocene sediments has often been interpreted to Globoconu sda ubjergen s(iBsronnimann).—Olsso 1n9,70:60 p91 fi2,.ig: . demonstrate the evolution of G. daubjergensis from G. 2a,b [lowe rDanian ,Pine Barren Mbr. ,Clayton Fm. ,Millers Ferry, cretacea (Olsson, 1970, 1982; Premoli Silva, 1977; Smit, 1977, Alabama ]f,ig s5.a-6 b[Danian b,asa Hl omerstow nFm N., e wJersey],— 1982; Li and Radford, 1991; Liu and Olsson, 1992). Keller 1, 993:1 p, i3. f:igs 1. - 3[Danian O, DP Hol e690C/15/1 1: 9-2 1cm], p i4. f:igs 1. 1-1 3[Danian H, ole 690C/15/1 1: 21-12 3cm W; edde lSl ea, Blow (1979:1235) suggested that G. daubjergensis, G. Sou Othceerann], daubjergensis gigantea ,and G .kozlowski ishould probably be Globasti cdaaubjergens (iBsronnimann).—Blo w19, 79:123 p57 i,f4.i:g 7s-.9 considered as only subspecifically related. Bang (1969) [Zon eP Di, SD PHol e47.2/11/2 1:48-15 c0m S;hatsk Ryis en,orthwestern propose dth esubspecie sdesignatio nGloboconus adaubjergen¬ Pacifi cOcean ]p, i2.56 f:igs 1.- 9[Zon eD2 D, anian K, arlstrup D, enmark], sis gigantea for morphotypes of G. daubjergensis that are p 2i.57 f:ig s3 4.[,uppe Dr anian O, stratorp S,weden], marked by an umbilical bulla and secondary bullae in Original Description. —“The specimens are very small intercamera lsutures .The holotype illustration of Globigerina for the genus [ Globigerina ]. The outline of the trochoid test is kozlowskii Brotzen and Pozaryska (1961) has a higher initial distinctly lobulate. The spiral side is pointed in the initial spire and tighter coiling mode than the holotype of G. portion. The umbilicus is small and shallow. The final whorl, daubjergensis .Nonetheless ,the former falls within the range of the dominant portion of the test, consists of 3 to 4 gradually in morphologic variability of the latter. Hence, G. kozlowskii is a size increasing subglobular chambers .The sutures of the final junior synonym o fG .daubjergensis. whor lare strongly incised ,those of the early test are not clearly Recrystallization renders the holotype o fPostrugoglobiger¬ visible. The extremely small, subcircular aperture opens into ina praedaubjergensis Salaj difficult to unambiguously iden¬ the shallow umbilica ldepression .The rough surface is covered tify (Salaj, 1986, pi. 3: fig. 7); however, Salaj (1986) by minute irregularly distributed spines. The thin walls are considered Postrugoglobigerin apraedaubjergensi simmedi¬ finely perforate. The direction of coiling is undetermined, 11 ately ancestral to G. daubjergensis, and the only other out of 19 specimens are coiling to the right hand side. illustrated specimen assigned to P. praedaubjergensis is a “The maximum diameter of the tests varies from 0.125 mm Globoconusa daubjergensis specimen (Salaj, 1986, pi. 3: figs. to 0.2 mm, average 0.15 mm. The height of the test is ±0.12 8 9, ). mm and the aperture has a diameter of ± 0.02 mm. Common in STABLE Isotopes. —Although its abundance in near-shore sample 38.” (Bronnimann ,1953:340.) sequences indicates a near-surface planktic habitat (Troelsen, 82 SMITHSONIA CNONTRIBUTION TS POALEOBIOLOGY FIGU R3 —E2.Paleobiogeograph mic asphowi ndgistributi oGonfloboconu sdaaubjergen s(iBsronniman inn) Zone sP an dPL 1957; Keller, 1989; Liu and Olsson, 1992), its oxygen isotopic Original Description. —“Very small species with 3-6 signature and open-marine abundance patterns suggest a chambers in the last-formed whorl; at the dorsal side all preference for relatively coo lwater masses (Premol iSilva and chambers visible; at the ventral side only those of the Boersma, 1989; D’Hondt and Keller, 1991; Liu and Olsson, last-formed coil ;chambers globular ,with sutures depressed on 1992 ;D’Hondt and Zachos ,1993). both sides; umbilica lcavity very small; walls thin ,consisting of Stratigraphic Range.—B asal Zone Pa to Zone Pic. one lamella ,with smal lpustules in between the pores ;often the Global Distribution. — Globoconusa daubjergensis was pustules are found in rows, as in Rugoglobigerina. Pores fine, abundan tin high-latitude and near-shore planktoni cforaminif- pipelike, at the surface ending in a funnel, as in some species eral assemblages (Troelsen, 1957; Keller, 1989; Premoli Silva they end in pits of the wall. Aperture an umbilical- and Boersma, 1989; Liu and Olsson, 1992). It was rare in extraumbilical ,crescent-like opening ,with slightly thickened low-latitude open-marine environments (Premoli Silva and border, or a narrow poreless lip.” (Hofker, 1978:60.) Boersma, 1989; D’Hondt and Keller, 1991; Liu and Olsson, Diagnostic Characters. —Original description inaccu¬ 1992 )(Figure 32). rate, especially with respect to surface texture. Test typically ORIGIN OF Species .— Globoconusa daubjergensis is consid¬ small, with moderate to flat trochospire consisting of 2'/2 ered to be a direct descendent of Guembelitria cretacea (Bang, 1969; Olsson, 1970, 1982; Premoli Silva, 1977; Smit, 1977, whorls of globular or subglobular chambers. Chambers 1982; Li and Radford, 1991; Liu and Olsson, 1992). gradually increase in size; 3 */2—7 chambers in each whorl, Repository. —Holotype (USNM CC64879) deposited in separated by radial and depressed sutures on both spiral and the Cushman Collection ,Nationa lMuseum of Natura lHistory. umbilical sides. Umbilicus closed. Aperture umbilical to Examined by SD, CL, and RKO. extraumbilical, ranging from comma-shaped arch to long narrow opening extending up apertura lface in nearly equatorial Genus Parvularugoglobigerina (Hofker ,1978) ,emended position. Aperture bordered by a slight lip. Surface texture microperforate ,pustulous or smooth; when pustulous ,well- Postrugoglobige Sr 1ian9la8j6, :52. preserved parvularugoglobigerinid sexhibi tpore-mound sur¬ TYPE Species. — Globigerina eugubina Luterbacher and face texture characteristic of guembelitriid taxa. Pores do not Premol iSilva ,1964. end in pits ;pustules not aligned. NUMBE 8R5 83 DISCUSSION. —As noted by several previous authors, Hof- ORIGIN OF Species.— This species evolved from Guembe¬ ker’s (1978) description of Parvularugoglobigerina eugubina litria cretacea .Parvularugoglobigerina alabamensis appears (Luterbacher and Premoli Silva) does not accurately describe as a morphologically intermediate form between G. cretacea Globigerina eugubina Luterbacher and Premoli Silva, 1964 and Parvularugoglobigerina extensa Blow. For example, the (Bang, 1979; Brinkhuis and Zachariasse, 1988; Loeblich and apertural characteristics of P. alabamensis more closely Tappan, 1988; Liu and Olsson, 1992; Olsson et al., 1992). resemble those of G. cretacea than do those of typical P. Nonetheless ,the genus Parvularugoglobigerina was erected extensa (compare Plates 63 and 65); however, the exact with Globigerina eugubina Luterbacher and Premoli Silva, phylogenetic relationship of Parvularugoglobigerina ala¬ 1964, as the type species, and it remains the generic label for bamensis to Parvularugoglobigerina extensa is unclear ,in part that species and related forms (Hofker, 1978; Brinkhuis and because the first known occurrence of P .extensa is stratigraph- Zachariasse, 1988; Loeblich and Tappan, 1988; Olsson et al., ically lower than the first known occurrence of P. alabamensis. 1992). It appears likely that either P. alabamensis evolved from G. The type species of Postrugoglobigerina is Postrugoglobig- cretacea independently of P. extensa (Liu and Olsson, 1992) or erina hariana Salaj, 1986. Because Postrugoglobigerina P. alabamensis was ancestral to P. extensa, but the known hariana Salaj, 1986, is a junior synonym of Globigerina stratigraphic record of P. alabamensis is incomplete. The first eugubina Luterbacher and Premol iSilva ,1964 ,Postrugoglo¬ hypothesis is stratigraphically simpler ,bu tevolutionarily more bigerina Salaj, 1986, is a junior synonym of Parvu¬ complex ,than the second because the forme rnecessarily entails larugoglobigerin aHofker 1, 978. iterative evolution o four-chambered parvularugoglobigerine Parvularugoglobigerinid gseneral lryesemb lwe oodringinids morphotype sfrom G .cretacea. in thei rsurface texture and apertura lcharacteristics. TYPE Locality. —Forty feet above the base of the Clayton Fm. at Millers Ferry, Alabama. Parvularugoglobigerina alabamensis (Liu and Olsson, 1992) Repository. —Holotype (USNM 460338) and paratypes(USNM 460339-460342) deposited in the Cushman Collec¬ Plate 65 :figures 1-6 tion, National Museum of Natural History. Examined by SD, Guembelitria allabamensi sLi uan dOlsson 1,992:341 p, i2 .f:igs 1.- 7[Zone CL, and RKO. Pla P,in eBarre nMbr C., layto nFm M., iller Fserr yA, labama]. Original Description. —“Test small, 110-160 pm in Parvularugoglobigerina eugubina (Luterbacher height and 90-135 pm in largest diameter, height-width ratio and Premoli Silva, 1964) 1.0-1.2; 10-13 globular or spherical chambers arranged in a Figure 33 ;Plate 66 :figures 1-12 ;Plate 67 :figures 1-14 high trochospire, triserial in the early and trochospiral in the later ontogenetic stage, with 3 to 4, mostly 3 V 2 to 4 chambers Globigerin anconitan Lauterbache arn dPremo Sliilv a1,964:10 7p ,2 if.:ig. in the last formed whorls; sutures deeply incised; umbilicus 3 aG -[clobigerin eaugubin Zaon eG,ubbi oce, ntr aAlppenine Ist,aly],Globigerin aeugubin aLuterbache arn dPremo lSiilva 1,964:105 p, i2 .f:ig. open ,narrow and shallow; aperture semicircular ,high ,with a 8a -[cGlobigerin eaugubin Zaon eG,ubbi oc,entr aAlppenine sIt,aly],— distinct narrow but thick lip ,umbilical ,symmetrically situated Premo lSi ilv a n dBoll i1, 973:526 p, i7. f:igs 2. - 5[Globigerin aeugubina at the base of the last formed chamber. Wall microperforate, Zone :figs .2-4 ,DSDP Site 152/10/1 :127-130 cm ;fig .5 ,DSDP Site early chambers have intensive small pore mounds, later 152/10/CC N; icaragu Rais eC,aribbea Snea],—Sm i1t,982:33 9p , 1if.:igs. chambers are covered with dense blunt pustules and less 1-20 p, i2. f:igs 1. - 8[Zon ePa G, redero s,outheaster nSpain].—Keller,1988:257 p, i .3 :figs .8-10 [Zone Pa E, Kl ef T, unisia] ;1989:321 p, i .6 :figs. frequen tpore-mounds.” (Liu and Olsson ,1992:341.) 3 6,[Zon ePa B, razo sRiver T, exas]. Diagnostic Characters. —Typically characterized by 2 V 2 Globigerin asabin aLuterbache arn dPremo lSiilva 1,964:108 p, i2 .f:ig 6.a-c whorls ,each composed of 3-4 inflated subglobular chambers, [Globigeri neaugubi nZaon Geu, bb icoe,nt rAapl penine Ista, ly]. increasing slowly in size. Aperture low, centrally located, Globigerin uambric Lauterbache arn dPremo Sliilv a1,964:10 6p ,2 if.:ig 2.a-c umbilical ,marked by a distinct lip .Sometimes characterized by [Globigeri neaugubi nZaon Geu, bb icoe,nt rAapl penine Ista, ly].Globorotali a(Turborotal i ca) fh.emisphaeric a(Morozova).—Blow, presence of pore-mounds (Plate 65 :Figure 6). 1979:1077 ,pi .64 :fig .8 [Zone Pa ,DSDP Hole 47.2/11/3 :148-150 cm; DISCUSSION —. Parvularugoglobigerin a labamensi sdiffers Shats Rkyi sneo, rthweste Prnac iOficcean], from Guembelitria cretacea by possessing 3 V 2 —4 (rather than Globorotal i Ta(urborotalia l)ongiapertur aBlow 1,979:1085 p, i5.6 f:ig s3 .4, 3) chambers in the outer whorl. It resembles G. cretacea in its [Zone Pa D, SDP Hole 47.2/11/6 :148-150 cm] p, i5. 8 :figs 3. -5 [Zone Pa, apertura lcharacteristic sand presence o fpore-mounds .Parvu¬ DSDP Hol e47.2/11/5 1: 48-15 0cm] p, i6. 3 f:igs 1. - 9[Zon ePa D, SDP Hole47.2/11/4 :148-150 cm] ,pi .68 :fig .3 [Zone Pa ,DSDP Hole 47.2/11/3: larugoglobigerina alabamensis sensu stricto differs from 148-150 cm] ,pi .72 :fig .1 [given as Zone PI ,DSDP Hole 47.2/11/1: Parvularugoglobigerina extensa in having a less elongate 148-15 c0m S;hats kRyis en,orthweste rPnacif Oiccean], aperture tha tdoe sno textend extraumbilically. Globorot aT l(uiarborota l c)ipa?fe. ntagon (aMorozova).—Blo w19, 79:1088, Stable Isotopes.—-N o data available. pi .64 :fig .1 [Zone Pa D, SDP Hole 47.2/11/4 :148-150 cm ;Shatsk yRise, Stratigraphic Range. —Zone Pa to Zone P3b. northwes tPear ncOifciecan].Globorota l(iTaurborotal ictaef).trago n(Ma orozova).—Blo 1w9,79:111 p3i,. Global Distribution.—S imilar to Parvularugoglobiger¬ 64 :fig .7 [Zone Pa ,DSDP Hole 47.2/11/3 :148-150 cm ;Shatsky Rise, i neaugubina. northwester nPacifi cOcean []i npar tn, o pt i6.7 f:ig 5.], 84 SMITHSONIA CNONTRIBUTION TS POALEOBIOLOGY Postrugoglobigerina hariana Salaj ,1986:53 ,pi .3 :figs .1-3 ,5 [base of Discussion. —Premoli Silva and Bolli (1973) declared Dania nHEl,ar iTau, nisia]. Globigerin ac fe.dit a(Subbotina).—Kelle r1,988:257 p, i3 .f:ig 1. 8[Zon ePa, Globigerina umbrica Luterbacher and Premol iSilva ,1964 ,and EKle Tf,unisia], Globigerina sabina Luterbacher and Premol iSilva ,1964 ,to be Globiger {inEaoglobigeri nhae)misphaer Micaorozova.—Kel l1e9r8, 8: 2p5i.7 junior synonyms of Globigerina eugubina Luterbacher and 3 f:igs 6. 7,1 1[Zon ePa E, Kl ef T, unisia] [,No tGlobigerin a{Eoglobiger¬ Premoli Silva, 1964. This broadened the concept of eugubina i nhae)misphaer iMcaorozo v1a9,61.] to include moderately spired forms with 4'/2 chambers in the Globigeri n{aEoglobigerin taa)uri cMaorozova.—Kelle 1r9, 88:25 pf73ii,g:. s. fina lwhor l(Globigerina sabina ) and forms with 7 chambers in 4 5[,Zon ePa E ,Kl e fT,unisia ][.No Gt lobigerin a{Eoglobigerina t)aurica Moro z1o9v5a7,.] the final whorl (Globigerina umbrica). We also include in this Parvularugoglobige eruingau b(Linuaterbache rParne dmSilovlai).—D’Hondt broadened concept Globigerina anconitana Luterbacher and and Keller ,1991:96 ,pi .4 :figs .4-6 [Zone Pa ,DSDP Site 577/12/5: Premoli Silva (1964), a species that was described from the 115-11 c7m S;hatsk Ryis en,orthwester Pnacif Oiccean],—L iaun Odlsson, same assemblage as these species .Smit (1982) first noted that 1992:345 ,pi .3 :figs .1-11 [Zone Pa ,Pine Barren Mbr. ,Clayton Fm., Globorotalia longiapertura Blow, 1979, is a junior synonym of Miller sFerry A, labama ][,No tHofker 1, 978:60 p, i2. f:igs 6. -10 p, i3. f:igs. 1 2,.] Globigerina eugubina Luterbacher and Premol iSilva. Parvularugoglobigeri nloangiapertu (rBalow).—D’Hon ad ntKdeller, Following Premoli Silva and Bolli (1973) and Smit (1982), 1991:96 ,pi .4 :figs .7 8 [Zone Pa ,DSDP Site 577/12/5 :115-117 cm; we treat several intergrading morphotypes as variants of P. Shats Rkyi sneo, rthweste Prnac iOficcean], eugubina (Plates 66, 67). These include a moderately spired Parvularugoglobigerin ma orphotyp 3De,’Hon datn Kdelle 1r,991:9 3p 3,i.: form with 4'/2-5 chambers (Plate 66: Figures 1, 2, Plate 67: figs .10 ,11 [Zone Pa ,DSDP Site 577/12/5 :115-1 17 cm ;Shatsky Rise, Figures 13, 14) in each whorl (= Globigerina sabina northwes tPear ncOifciecan]. Parvularugoglobigeri necfua.gubi n(Lauterbach ae nPrdrem Soilliva).—Liu Luterbacher and Premoli Silva, 1964; Globorotalia (Turboro- and Olsson 1, 992:345 p, i3. f:ig 1. 2[Zon ePa P, in eBarren Mbr .C, layton talia) longiapertura paratypes of Blow ,1979 ,Plate 63: Figures Fm M.,ille rFserr Ay,labama], 1-3; Parvularugoglobigerina morphotype 3 of D’Hondt and Keller, 1991; Parvularugoglobigerina cf. eugubina of Liu and ORIGINAL Description. —“Test very small, trochospiral; Olsson ,1992 )and a relatively low-spired form characterized by umbilical side flat, spiral side almost flat; the initial part of the 5-8 chambers per whorl (Plate 66: Figures 7, 8, 10-12, Plate spire is slightly raised relative to the final whorl. Composed of 67: Figures 1,3 ,10 ,12) that has an apertura lextension from the 12 to 14 globular chambers, disposed in 2 x h whorls; the final umbilicus to a near peripheral location and slightly more whor lcontains 6 chambers that increase gradually in size .The embracing chambers with less deeply incised sutures (the fina lchamber occupies about ‘/4 to Vs of the test’s surface .The longiapertura morphotype o fBlow ,1979). periphery is lobate. Sutures depressed and radial on the Globigerina minutula Luterbacher and Premol iSilva ,1964, umbilica lside ,depressed and slightly arcuate on the spira lside. was described along with Globigerina eugubina from the Umbilicus quite large .Aperture at the base of the fina lchamber Gubbio section. The holotype drawing of G. minutula in an umbilica lposition .Surface slightly rugose. illustrates a very low trochospiral, 3-chambered form, which Dimensions: closely resembles Plate 65 :Figure 13 that was selected from the type level. The specimen is completely recrystallized so that it is uncertain whether it is a microperforate or norma lperforate taxon. Consequently, given its generalized morphology, we canno taccurately identify this taxon. Although badly preserved ,the type specimen o fPostrugog¬ lobigerina hariana Salaj, 1986, is indistinguishable from Parvularugoglobigerina eugubina .The holotype o fP .hariana (Luterbacher and Premoli Silva, 1964:105; translated from is a low trochospira lform with a pore-mound surface texture, Italian.) five globular chambers in each of two whorls, a low rate of Diagnostic Characters. —Test a small trochospire with chamber expansion, and a “narrow slot-like interiomarginal 2 '/2 whorls, each whorl with 4'/2—8 inflated subglobular umbilical aperture at the base of the last chamber” (Salaj, chambers; chambers increasing slowly in size; first and last 1986:53) .Consequently ,Postrugoglobigerina hariana Salaj, whorls usually with same number of chambers. Spire height 1986 ,is regarded as a junior synonym o fParvularugoglobiger¬ low to moderate. Test wall microperforate, well-preserved ina eugubina (Luterbacher and Premoli Silva, 1964). In specimens sometimes exhibi tsurficia lpore-mounds (Plate 67: addition, we include in P. eugubina a number of forms Figure 14). Aperture elongate, marked by thickened rim; identified by Blow (1979) and Keller (1988) to various species apertura lposition ranges from umbilica lto nearly peripheral of cancellate Danian species (see synonomy) .The illustrations (Plates 66 ,67) .Individua lspecimens sometimes exhibit dorsal of Blow and Keller clearly show a microperforate wall aperture (Plate 66: Figure 6) or both dorsal and umbilical structure; therefore, these identifications are incorrect. The apertures on fina lchamber (Plate 66: Figure 9); some showing general morphology of their illustrated forms falls within the more than one dorsa laperture. broadened concep to fP .eugubina. NUMBE 8R5 85 FIGU 3R—3E.Paleobiogeograp hmi csahpowi ndgistribut Pi oanfrvularugoglobiger ienuagub i(nLauterbacher an dPremo lSi ilva i) nZon ePa. Stable Isotopes. —Oxygen isotopic data suggest that P. Eoglobigerina flodina Blow 1, 979:1221 p, i5. 7 :figs 5. 6 ,[Zone Pa D, SDP eugubina occupied a slightly deeper or cooler-season planktic Ho l4e7.2/11/ 51:48-15 c0m S;hatsk Ryis en,orthwester Pnacif Oiccean]. niche than co-occurring G. cretacea and Woodringina spp. Globigerin ma inutu lLauterbach earn Pdremo Slilva.—Sm i1t,982:33 8p 3,i.:fig s3.- 6[Zon ePa G, redero s,outheaster nSpain ][,No Ltuterbache arnd (Boersma et al., 1979; Boersma, 1984a; D’Hondt and Zachos, Prem oSliilv 1a9,64.] 1993). Globoconus caonus (aKhalilov).—Kelle r1,988:25 7p , 3if.:ig s1.2-1 [4Zone Stratigraphic Range. —Zone Pa. POb ,E lKef ,Tunisia] ;1989:319 :figs .3 ,9-11 [Zone Pa ,Brazos River, Global Distribution. —Biogeographic data indicate that Texa s[]N. oKthalilo 1v9, 56.] P. eugubina was a low to middle latitude taxon with an Parvularugoglobigerin ma orphotyp 1De,’Hond atn Kdelle 1r,991:9 3p ,3i.: open-ocean affinity (Premoli Silva and Bolli, 1973; Premoli figs .3-5 ,7-10 [figs .3 ,4 ,7-10 ,Zone Pa ,DSDP Site 577/12/5 :115-117 Silva and Boersma, 1989; D’Hondt, 1991; D’Hondt and Keller, cm S;hatsk Ryis en,orthwester nPacif iOc cean f;ig 5l.o, we Dr anian B,razosR iTverx,as], 1991; Liu and Olsson, 1992) (Figure 33). Parvularugoglobigerin ma orphotyp 2De,’Hon datn Kdelle 1r,991:9 6p 4,i.1: Origin of Species. —This species evolved from Guembe- figs .1 ,2 [Zone Pa ,DSDP Site 577/12/5 :115-117 cm ;Shatsky Rise, litria cretacea via Parvularugoglobigerina extensa at the base northwe tPeam cOifcicean]. o fZone Pa. Parvularugoglobigeri na efafu.gubi n(Lauterbach ae nPrdrem Soilliva).—Liu Repository. —Holotype (No. C20532) and paratype (Nos. an dOlsson 1, 992:345 p, i2. f:igs 8. -1 1[Zon ePa P, in eBarre nMbr .C, laytonFm M.,ille Frserr Ayl,abama]. C20554, C20555) are deposited in the Basel Museum of Natura Hl istory S, witzerland. Original Description. —“The small test is coiled in a distinct, tightly expressed, trochospire with 9-10 chambers Parvularugoglobigerina extensa (Blow ,1979) comprising the spire and four chambers in the last whorl. In dorsal aspect, the chambers are inflated, closely appressed, Plate 65 :figure s7-13 closely-set and quite strongly embracing and the dorsal Woodringin haornerstownens gisrou Ppr,em oSliil vaan Bdo l1li9, 73:53 584, 0, intercamera lsutures are subradially to sinuously disposed .The p i6. f:ig 1. 0 p, i7. f:ig 1.[Globigerin aeugubin aZone D, SD PSit e150/10/2: 78-5 c0m C;aribbea nSea ][,No Ot lsson I,960.] dorsal and ventral intercameral sutures are depressed to Eoglobigerina el xtens aBlow 1, 979:1220 p, i6. 9 f:ig 7.[Zon eP ID, SD PHole moderately incised .In ventra laspect ,the chambers are inflated, 47.2/11/3 :0-5 cm] ,pi .74 :figs .1 ,2 [Zone PI ,DSDP Hole 47.2/11/3: subglobular, but embracing. The umbilicus is closed and the 148-15 c0m S;hats kRyis ne,orthweste rPnaci fOiccean]. aperture extends from the umbilicus extensively towards the 86 SMITHSONIA CNONTRIBUTION TS POALEOBIOLOGY anterior side of the last chamber and is a widely open, almost maintained in future studies .On the basis of wal lstructure and rectangular, opening bordered by a weakly developed but texture, Blow (1979) suggested that extensa and fodina (= thickened porticus. The equatorial profile is strongly lobulate extensa ) were more closely related to Globastica (= Globocon- but almost circular in outline whilst the axial profile shows the usa ) daubjergensis than to any other species assigned to distinct early trochospire and subglobular nature of the later Eoglobigerina .He further hypothesized that “either fodina or chambers. The wall is very finely perforate and the mural-pits extensa migh tbe ancestra lto daubjergensis or ,alternatively the are very widely and sparsely developed ;the mura lpores do not various forms only share a common ancestor” (Blow, open into pore pits but the wall bears fine pustules which may 1979:1221). The latter interpretation is consistent with the be the bases of norma lspines .Maximum diameter of holotype presen thypothesi stha tGloboconusa daubjergensi sand Parvu¬ 0.198 mm. as measured electronically. The name extensa larugoglobigerina extensa both evolved directly from Guem¬ derives adjectively from the extensive nature of the aperture.” belitria cretacea (Olsson, 1970, 1982; Premoli Silva, 1977; (Blow ,1979:1220.) Smit, 1982; Liu and Olsson, 1992; Olsson et al., 1992). Diagnostic Characters. —Characterized by 2‘/2 whorls, Stable Isotopes. —No data available. each composed of 3 V 2 —4 inflated subglobular chambers, increasing slowly in size. Aperture centrally located, umbili- Stratigraphic Range. —Upper Zone P0 through Zone Pa. cal-extraumbilical ,marked by distinct lip ,often elongate ,and GLOBAL Distribution. —Similar to Parvularugoglobiger¬ often asymmetrically oriented. Test wall microperforate. i neaugubina. Despite origina ldescription ,pustules are not bases of normal ORIGIN OF Species. —This species evolved from Guembe¬ spines bu tguembelitriid pustules typically either perforated by litr icaretacea. pores (pore-mounds )o rperipherally associated with pores. Repository. —Holotype (BP Cat. No. 39/60) and paratype DISCUSSION .— Parvularugoglobigerina extensa i sa moder¬ (BP Cat. No. 40/24) are deposited at The Natural History ately high-spired form and is morphologically intermediate Museum ,London .Examined by GJ. between Guembelitria cretacea and Parvularugoglobigerina eugubina .Parvularugoglobigerina extensa primarily differs Genus Woodringina Loeblich and Tappan, 1957 from G. cretacea sensu stricto by possessing 3 V 2 —4 (rather than 3) chambers in the outer whorl and by typically having an Type Species. — Woodringina claytonensis Loeblich and elongat eapertur etha et xtend sextraumbilically. Tappa 1n9, 57b. Eoglobigerina ̂fodina Blow ,1979 ,was originally described Original Description. —“Test free, early stage with a as characterized by a small but open umbilicus and a circular single whorl of three chambers, followed by a biserial stage; aperture “extending from the opening in a slightly oblique chambers inflated; wall calcareous, radial in structure, finely manner, into the terminal face of the last chamber” (Blow, perforate ;aperture a low arched slit ,bordered above by a slight 1979:1221) .The name fodina was derived from the Latin word lip .Family Heterohelicidae .Paleocene .Monotypic.” (Loeblich for pit or tunnel and applied to the apertural shape, which was an dTappan 1, 957b:39.) described as looking like the entrance to a London Under¬ Diagnostic Characters. —Tests contain an initial whorl ground Railway tunnel (Blow, 1979:1221). The slight differ¬ of 3 chambers, later whorls of 2 chambers each. Test wall ences between the holotypes of extensa and fodina result from microperforate, marked by a guembelitriid surface texture elongation of the former’s aperture toward the umbilicus. (smooth walled o rbearing perforate pustules) .Aperture usually Given the extreme plasticity of apertural morphology and asymmetrically positioned and thin apertural lip infolded on location in related taxa (i.e. ,Parvularugoglobigerina eugubina on seide. and Woodringina species), such minor differences appear Discussion. —Although Woodringina remains limited to insufficient to maintain extensa and fodina as separate taxa. the Paleocene ,it is no longer monotypic .It was assigned to the Consequently ,we have reduced Eoglobigerina! fodina Blow, subfamily Guembelitriinae by Loeblich and Tappan (1957b). 1979 ,to a junior synonym of Parvularugoglobigerina extensa Elevation of the subfamily Guembelitriinae to the family (Blow, 1979). In addition, the 3'/2-4 chambered inflated Guembelitriidae (El-Naggar, 1971; Blow, 1979; Loeblich and subglobular ,microperforate forms described by Premol iSilva Tappan ,1988) removed it from the family Heterohelicidae. and Bolli (1973), Smit (1982), Keller (1988), D’Hondt and Keller (1991), and Liu and Olsson (1992) under various names Woodringina claytonensis Loeblich and Tappan, 1957 (see synonomy) are placed in P .extensa. Although Blow (1979:1220) provisionally placed extensa Plate 13 :figures 6-8 ;Plate 68 :figures 1-6 and fodina in the genus Eoglobigerina, he recognized their Woodringin calaytonens iLsoeblic ahn Tdappa n1,957b:3 9fi: gla. - [dlower microperforate structure as quite “distinct from the normal Danian P, in eBarre nMbr .C, layto nFm .A, labama ]1;957a 1:78 p, i4.0 f:ig 6. perforate, cancellate or reticulate, wall of the eobulloides- [holotyp ereillustrated].—D’Hond t1,991:172 p, i1 .f:ig s8 .1,1 1, 2[fig s8.,12 D, SDP Sit e577/12/5 9: 4-9 6cm f;ig 1. 1 l,owe rDanian B, razo sRiver, trivialis-edita group.” On that basis, he expressed doubt that Texas] ,pi .2 :figs .4 ,12 [DSDP Site 577/12/5 :94-96 cm ;Shatsky Rise, assignment of those forms to Eoglobigerina would be northwester nPacif iOc cean],—Li uan dOlsson 1,992:341 p, i1 .f:ig s4.-6 NUMBE 8R5 87 [Zone Pa P, in eBarren Mbr .C, layton Fm .M, iller sFerry A, labama],— separate taxon, given the pronounced variability in position, MacLeod 1, 993:61 p, i3. :figs 8. -14 [ODP Hole 690C/15X/2 :28-30 cm; Ma uRdis Seo, uthe rOncean], height ,and number of apertures exhibited by otherwise similar Woodringin haornerstownens Oislsson.—Kelle 1r9, 88:25 p 7f3,i :.g 1[5.Zone specimens in populations of Woodringina and other guembe- Pa ,E lKef ,Tunisia] ;1989:319 :figs .3-6 [lowe rDanian ,Brazos River, litriid genera (i.e. ,Guembelitria and Parvularugoglobigerina). Texa [sN] .Wotoodringi nhaornerstownen sOislsso In96, 0.] Hence, we consider Woodringina kelleri MacLeod, 1993, a Woodringin akelle rMi acLeod 1,993:63 p, i4. f:igs 1. - 3[Danian D, SD PSite junior synonym of Woodringina claytonensis Loeblich and 577A/12/ 24:4-4 c6m S;hatsk Ryis en,orthweste rPnacif Oiccean], Woodringina cf .keller iMacLeod 1, 993:63 p, i .3 :figs .4 5, 1, 2 [Zone Pa E, l Tappa n1,957. KTeufn, isia], Stable Isotopes. —The stable isotopic signature of this species suggests an affinity for relatively warm near-surface Original Description. —“Test free, tiny, flaring rapidly; water masses (D’Hondt and Zachos ,1993). early stage with a single whorl of three chambers (reduced Stratigraphic Range. —Basal Zone Pa to Zone Plb. ‘triserial’), commonly followed by three or more, rarely up to five, pairs of biserial chambers, the plane of biseriality slightly GLOBAL Distribution. — Woodringina claytonensis was twisted in development ;chambers few in number ,subglobular, most abundant in low-latitude open-ocean assemblages increasing rapidly in size; sutures distinct, constricted; wall (D’Hondt and Keller, 1991; Liu and Olsson, 1992). It was rare calcareous, finely perforate and very finely hispid; aperture a in high-latitude open-marine environments (Liu and Olsson, low arched slit bordered above by a slight lip, somewhat 1992). asymmetrica i lnposition. Origin of Species. —This species is considered to have “Length of holotype 0.15 mm.; greatest breadth 0.12 mm. descended from Guembelitria cretacea (Olsson, 1970, 1982; Other specimens range from 0.12 to 0.22 mm. in length.” Smit, 1977, 1982; D’Hondt, 1991; Li and Radford, 1991; (Loeblich and Tappan ,1957b:39.) Olsson etal., 1992; Liu and Olsson, 1992). D’Hondt (1991) and Diagnostic Characters. —In general, original description Liu and Olsson (1992) illustrated forms that are morphologi¬ accurately describes diagnostic features of W. claytonensis, cally intermediate between Guembelitria cretacea Cushman, except for description of microperforate wall as hispid. Some 1933, and Woodringina claytonensis Loeblich and Tappan, specimens smooth-walled (Loeblich and Tappan ,1988) ,others 1957. Such morphotypes are triserial in early whorls and characterized by presence o fscattered pore-mounds (Plate 68: biserial in later whorls ( Guembelitria irregularis Morozova, Figure 6). In other respects, species highly variable 1961, of D’Hondt, 1991) (Plate 63: Figure 12; Plate 68: Figure morphologically ,with apertura lheigh tvaring from high to low. 5). Twisted plane o fbiseriality ,apertura lasymmetry ,and triserial- Repository. —Holotype (USNM P5685) deposited in the ity of the first whorl more strongly pronounced in some Cushman Collection, National Museum of Natural History. specimens than in others within same assemblages. Examined by SD, CL, and RKO. DISCUSSION .—MacLeod (1993:92 )described Woodringina keller ias distinguished from Woodringina claytonensis by the Woodringina hornerstownensis Olsson ,1960 former’s “laterally compressed adul tchambers ,and .. .its large elongate aperture that is surrounded by a well-developed Figure 34 ;Plate 13 :figures 4 ,5 ;Plate 68 :figures 7-14 discontinuous apertura lrim;” however ,the fina lchambers of Woodringin haornerstownens iOs lsso n1,960:2 9p , 4if.:ig s1. 81 ,[9Zon Pe3b, both holotypes are similarly subglobular in edge and plan view Homerstow nFm N., e wJersey],—D’Hond t1,991:17 2p, 2i .f:ig s5.- [8figs. (compare MacLeod, 1993, pi. IV: figs. 1-3 to Loeblich and 5 8 ,Z,on ePla D, SD PSit e577/12/4 3:4-3 6cm S;hatsk yRise n, orthwesternPacifi cOcean f;igs 6 .7 ,Z, on eP3b H, omerstow nFm .N, ew Jersey].—Liu Tappan, 1957b, pi. 39: fig. 1; see also Plate 13: Figures 6, 7). and Olsson 1, 992:341 p, i .1 :fig .8 [Midway Group P, lumme rStop 14, The only other figured specimen identified by MacLeod (1993) Kaufman Co .T, exas].—MacLeod 1, 993:63 p, i4. f:igs 6. 7, 1, 1 1, 3[lower as W. kelleri also lacks laterally compressed chambers and Dania n EK,le Tf,unisia], closely resembles the W claytonensis holotype in the shape of Chiloguembelitr itaauric Ma orozova.—Kelle r1,988:25 7p , 3if.:ig [3Z. one its final chambers (compare D’Hondt, 1991, pi. 2: fig. 4 to P a EK,le Tf,unisia []N. o Cthiloguembelitr itaauric Ma orozov a1,961.] Loeblich and Tappan, 1957b, pi. 39: fig. 1). Comparison of the Original Description. —“Test small, elongate, slightly respective apertural characteristics of W claytonensis and W twisted, rather rapidly tapering, about twice as long as broad. kelleri is complicated by the absence of W claytonensis Initial end consists of whorl of three chambers, rest of test paratype specimens and the physical obstruction of the W consists of biserial arrangement of chambers. Wall smooth, claytonensi sholotype aperture .Nonetheless ,the W claytonen¬ surface of last two chambers may be finely spinose, especially sis holotype also appears to have a large elongate aperture near the aperture. Chambers numerous, somewhat inflated, surrounded by a well-developed ,discontinuous apertura lrim wider than high ,increasing more rapidly in width than height as (Plate 13: Figure 7). The aperture of the W. kelleri holotype added. Sutures distinctly depressed, nearly at right angles to (Plate 13: Figure 8) does appear to be more centrally located axis of test. Aperture a low arch at one side of final chamber, than that of the W. claytonensis holotype. This difference directed inward ,with a smal llatera lflange along the periphery. appears insufficient to warrant maintenance of W kelleri as a Length 0.15 mm. Width 0.10 to 0.15 mm.” (Olsson, 1960:29.) 88 SMITHSONIA CNONTRIBUTION TS POALEOBIOLOGY Diagnostic Characters. —Test wall microperforate; aper¬ Stable Isotopes. —Biogeographic and stable isotopic data ture marked by a thin lip. Apertural position and height suggest an open-ocean, warm shallow-water habitat for W. somewhat variable. Biserial portion of test distinctly twisted, hornerstownensis ,similar to that of W .claytonensis (D’Hondt although less so than W claytonensis. Woodringina horner- and Keller, 1991; Liu and Olsson, 1992; D’Hondt and Zachos, stownensis distinguished from W. claytonensis by “the elon¬ 199 3. ) gate tapering test and the almost straight sutures” (Olsson, Stratigraphic Range. —Zone Pa to Zone P3b. 1960:29). Woodringina hornerstownensis often with six or Global Distribution. —Widespread in high and low more pairs of biserial chambers, whereas W claytonensis latitude s(Figur e34). usually limited to five or fewer. Origin of Species. —This species is generally considered to DISCUSSION .—The holotype illustration and description of have descended from Guembelitria cretacea via W .claytonen¬ Chiloguembelina taurica Morozova ,1961 ,closely resembles sis (Olsson, 1970, 1982; Smit, 1977, 1982; D’Hondt, 1991; Li W hornerstownsis Olsson ,1960 .Chiloguembelina taurica was and Radford, 1991; Olsson et al., 1992; Liu and Olsson, 1992). originally described as being characterized by a “high narrow Repository. —Holotype (USNM 626457) deposited in the test, weakly compressed bilaterally, its height two to three Cushman Collection, National Museum of Natural History. times the width. Lateral outlines at first subtriangular, later Unfigured paratypes deposited at Princeton University (No. almost parallel” (Morozova, 1961:18). The test of C. taurica is 81038) and Rutgers University (No. 5026). All specimens formed of 10-12 spheroidal chambers, and its intercameral examined by SD, CL, and RKO. sutures are almos tstraight .Although no tvisible in the holotype illustrations, the aperture of C. taurica was described as semicircular and basal (Morozova, 1961). Based on its type Family Chiloguembelinidae Reiss, 1963 illustration and description, Chiloguembelina taurica Mo¬ rozova ,1961 ,should be considered a possible junior synonym (by S. D’Hondt and B.T. Huber) o fWoodringina hornerstownensis Olsson ,1960 .The holotype Original Description. —“Trochospirally coiled with two illustrations and descriptions o fHeterohelix gradata Khalilov, chambers per coil arranged around an elongated axis. Test 1967, and Heterohelix gradata normalis Khalilov, 1967, also usuall ytwisted A. perture single a, n interiomargina al symmetri¬ closel yresemble Woodringina hornerstownensi sOlsson 1, 961. cal low to high arch, bordered by an asymmetrically situated Heterohelix gradata normalis was originally described as being characterized by an elongate “wedge-shaped [test], flap which is often protruding and platelike, or terminal, graduall ybroadening toward th eobliquel ytrimmed-of fapertu¬ situated on a shor tneck .No distinc ttoothplates .Ornamentation ra lend .. .chambers biserially arranged ,in each offset row there if present consisting of inflational papillae or short spines.” are 6-8 spherical chambers. ... Aperture semilunate, much (Rues s1,963:55.) shifted from the median frontal position and shielded on one Diagnostic Characters. —Small test comprised of biseri¬ side by a moderately protruding lip” (Khalilov, 1967:173). ally arranged chambers, often with a slightly twisted coiling From this description H .gradata normalis is indistinguishable axis. Intercameral sutures distinct, depressed, and often from W .hornerstownensi sin genera ltes tmorphology .Further¬ somewhat oblique .Wal lcalcareous and microperforate with more ,it appears to share the asymmetry o fapertura lshape and smooth to pustulous surface texture .Aperture arched ,rimmed position that is diagnostic of Woodringina and related taxa. by narrow lip, and generally infolded on one side of ultimate Heterohelix gradata sensu stricto was distinguished from H. chamber .Many Chiloguembelina species with well-developed gradata normalis by its last 4-10 chambers being much larger flap or flange bordering aperture along infolded side. than the preceding chambers (Khalilov ,1967). It also appears Discussion. —As noted in the discussion of the Guembe- to share the chamber shape ,adult chamber arrangement ,and litriidae ,the Chiloguembelinidae appears to be a monophyletic apertural asymmetry of W. hornerstownensis. Given the or paraphyletic family descended from Guembelitria cretacea genera lcongruence of their origina lillustrations and descrip¬ via Woodringina claytonensis (Olsson, 1970, 1982; Premoli tions with W .hornerstownensis ,Heterohelix gradata Khalilov Silva, 1977; Smit, 1982; D’Hondt, 1991; Li and Radford, 1991; and H. gradata normalis Khalilov should be considered Liu and Olsson, 1992; Olsson et al., 1992). It is generally possible junior synonyms of W. hornerstownensis Olsson, believed that this family was monophyletic throughout the 1960. The original descriptions of H. gradata and H. gradata Paleocene. It’s post-Paleocene status is less clear, as the normalis stated that the tests of these taxa are covered with relationships of Chiloguembelina to such late Paleogene and large pores (Khalilov, 1967). If so, these taxa are distinguish¬ Neogene taxa as Streptochilus Bronnimann and Resig, 1971, able from W. hornerstownensis on the basis of wall structure. and Cassigerinella Pokorny ,1955 ,is presently uncertain. Close examination of wall structure and other relevant Reiss (1963) originally assigned both Chiloguembelina and characters (i.e. ,the presence or absence of initia ltriseriality) in Zeauvigerina to the Chiloguembelinidae. Given the Late type populations of H. gradata and H. gradata normalis are Cretaceous occurrence of Zeauvigerina (Huber and Boersma, necessary to conclusively define the taxonomic status of these 1994 )and the earlies tPaleocene derivation o fChiloguembelina taxa relative to W .hornerstownensis. from a guembelitriid ancestor, retention of this assignment 122 SMITHSONIA CNONTRIBUTION T SPOALEOBIOLOGY PLATE 8 Russian Primar yType Specimens (bar =s10 0pm) FIGURE S1-3.— Guembelitri airregular iMs orozova 1,961:17 p, i1 .f:ig 9 .h,olotyp eno 3.510/13a M, oscow GA ND;ania Tna, rkhanku Ctr, ime Sa e.“eDiscussio nf o”Gruembelitr ciaretacea. FIGURE S4-6.— Globigerin a(Eoglobigerina t)rifoli aMorozova 1,961:12 p, i1 .f:ig 1 .h,olotyp eno 3.510/4, Mosco GwA ND;ania Tna, rkhank uCtr,ime Sa e“. eDiscussio nf oG”rloboconu sdaaubjergensis. FIGURE 7S-9.— Acarinin ma ultiloculat Ma orozov a1,961:1 5p ,2 if.:ig 5h.,olotyp neo 3.510/1 0M, osco wGAN; Montia Bn,al kNaasypkoiskay Ca,rime Pa.roba br aelyworke sdpecime oHnfedberge lplalanispi r(aTappan, 1940). FIGURE 1S0-12. —Globigerin (aEoglobiger ien )oabulloid eMsorozov a19, 59:111 t5e,xt-fi gla.- hc,olotyp neo. 3508/ M1,osco GwA ND;ania Tna, rkhanku Ctr,ime Sa e.Eeoglobigeri neaobulloides. FIGURE 1S3-15.— Globigerin E (aoglobigerina h)emisphaeric Ma orozov a1,961:1 1p , 1if.:ig 4h.,olotyp neo. 3510/ M3,osco GwA ND;ania nTa, rkhanku Ct,rime aSe. “eDiscussion f ”oEroglobigerin eadita. FIGURE S16-18.— Globigerin aE(oglobigerin at)heodosic aMorozova 1,961:11 p, i1 .f:ig 6 .h, olotyp eno. 3510/ M2,osco GwA ND;ania nTa, rkhanku Ct,rime aSe. “eDiscussion f ”oEroglobigerin eadita. NUMB E8R5 123 O. &W 130 SMITHSONIA CNONTRIBUTION T SPOALEOBIOLOGY PLATE 12 Russian Primar yType Specimens (bar 1=s0 (0am) FIGURE S1-3.— Acarinin acarinat aSubbotina 1,953:229 p, i2.2 f:ig 5 .p, aratyp eno 4.129 S, tP. etersburg VNIGR (1378/160 z);on o ecfompresse gdloborotaliid sP,aleocene-lowe Erocen eS,erie F sKI,hie Ruiver, Caucasu Ss e.“eDiscussio nf oA”rcarini nnaitida. FIGURE S4-6.— Acarinin aintermedi aSubbotina 1,953:227 p, 2i.0 f:ig 1.5 h,olotyp eno 4.095 S, tP.etersburg VNIG R(3I78/124 z)o; n oceof mpresse gdloborotaliid Ps,aleocene G?,oryach Kelyn chhorizo nK,uba Rniver, Caucasu Ss e.“eDiscussio nf oA”rcarini nnaitida. FIGURE 7S-9.— Guembelitr idaammu lVaoloshin a1,96 1h,ypotyp eP;aleocen eP, 0B,ja l(aK/ sTectio n o2.b, 1 -c2 mabo vbeoundary B),ulgar iSae. “eDiscussion f ”oGruembelitr ciaretacea. FIGURE 1S0-12. —Globanomalin ima ita t(aSubbotin 1a9, 53 h),ypotyp eP;aleocen Pe,lb/ Bc,ja (laK /sTectio nno. 2 bsa, mp Sleu m24/12 B),ulgar iSae. Gelobanomalin ima itata. NUMB E8R5 131 132 SMITHSONIA CNONTRIBUTION T SPOALEOBIOLOGY PLATE 13 USNM Primary Type Specimens (bar s=5 0pm) FIGUR 1E.— Rectogiimbelin caretace Caushma n1,93 2h,olotyp eU,SN MCC1630 8u;ppe Mr aastrichtian, Arkadelp hCial aHyo, pAer,kansas. FIGUR 2E.— Tubitextular ilaeviga tLaoeblic ahn Tdappa n1,95 (7R=ectoguembelin caretace Caushman), holotyp eU,SN MP582 0lo; we Praleocen eM, cBryd Leimeston Me b rC„layto Fnm W., ilco Cxo A.,labama. FIGUR 3E. —Guembelitr icaretace Caushma n1,93 3h,olotyp eU,SN MC 1C902 2u;pp eMraastrichtia nN,avarro F Tme.x,as. FIGUR E54S., —fVoodringin haomerstownens Oislsso n19, 6 h0,olotyp Ue,SN M62645 Z7;on Pe3 bH,omerstown Fm N.,e Jwersey. FIGURE 67S,.— Woodringin calaytonens iLsoeblic ahn Tdappa n1,95 7h,olotyp eU,SN MP568 5lo; we Dranian, Pin Bearre Mn b rC.,layto Fnm A.,labama. FIGUR 8E.— Woodringin kaelle Mri acLeo d1,99 (3W= oodringin calaytonens iLsoeblic ahn Tdappan Z);on Pea, DSD SPi t5e77A/12/ 24:4-4 c6m S;hatsk Ryis en,orthwester Pnacif iOccean. FIGURE 9S1,0.— Gumbelin ma idwayens iCsushman 1,94 0h,olotyp eU, SNM CC3571 5b;as aMl idwa Fym., Sum tC eAorl.a, bama. FIGURE 1S 1,6.— Gumbelin tarinitatens iCsushma ann Rden z1,94 2h,olotyp eU,SN MCC3819 8P;aleocene, Solda Fd moTr.i,nidad. FIGURE 1S 21,3.— Chiloguembelin ma idwayens isstrombiform iBseckman n1,95 (C7=hiloguembelina midwayen s(iCsushman )h)o, lotyp Ue,SN PM577 G1;loborota lpiaseudomenar dZioi n Lei,za rSdprin gFsm., Trinidad. FIGURE 1S 41,5.— Giimbelin amorse Kiline 1,94 3h,olotype U, SNM 487301 D; anian P,orter Csree Ckla yC,lay MCois.s, issippi. FIGUR E1 S178, . —Chiloguembelin saubtriangular Biseckman n19, 5 h7,olotyp Ue,SN MP578 G3;loborotalia pusil lpausil lZaon elo, we Lrizar Sdpring Fsm T.,rinidad. Figure 1s 92,0.— Giimbelin awilcoxens iCsushma nan dPonton 1,93 2h,olotype U, SNM 16218 W; ilco Fxm., O zAalarkb,ama. NUMB E8R5 133 136 SMITHSONIA CNONTRIBUTION T SPOALEOBIOLOGY PLATE 15 USNM Primary Type Specimens (bar s=5 0pm) Figure 1s-3.— Globorotal isatrabocel lLaoeblic ahn Tdappa n1,95 7h,olotyp eU,SNM P587 9N;anafal iFam., Alabama. FIGURE 4 S78,,.— Globigerin saoldadoens iBsronniman n1,95 2h,olotyp eU,SN M37008 5L;izar Sdpring Fsm., Trinidad t;yp elocalit yo Gf loborotali avelascoens iZson eo Bf oll i1,957 a=Zon eP5. FIGURE 5S6,.— Globigerin aquiens iLsoeblic han dTappa nh,olotyp eU,SNM P583 9A;qu iFam A.,qu iCareek, Virginia. FIGURE 91S,0. —Globigerin cahascanon Laoeblic ahn Tdappa n1,95 (A7=carinin saubsphaeri c(aSubbotina)), holotyp eU,SN MP584 2Z;on Pe 4u,ppermo sHtomerstow Fnm N.,e wJersey. FIGURE 1S 1, 21,5.— Globorotal icarassat (aCushman v)a ra.equ Caushma ann Rden z1,94 2h,olotyp eU,SNM CC3821 0n;e abras oeGfloborotal isaubbotina Zeon eS,oldad Fom T.,rinidad. FIGUR E1 S134, . —Globigerin daaubjergens Bisronniman 1n9, 5 h3o, lotyp Ue,SN MCC6487 D9;ania Dn,aubjerg Qu Daernrym, ark. NUMB E8R5 137 mm 232 SMITHSONIA CNONTRIBUTION TS POALEOBIOLOGY PLATE 63 Guembelitri acretace aCushman 1, 933 (Figure s1-5 7, -12 :bar s= 50 jam ;Figure 6 :ba r= 10 (am) FIGUR E1S-4.—Upp eMraastrichtia Rne, dba nFkm N.,e Jwersey. FIGURE S5 6, 1, 0-12.—Zon ePa M, iller sFerry A, labama c,or e226 s,ampl e18 F; igur e6 v, iew o wf a lol Ff igure s5howin gtypica plor emound os tfh ispecie sF;igur e1 2s,pecime nshowin gtransitiona ml orpholog tyo Woodringina. FIGUR 7E.—Zon Pe aD,SD HPo l3e90A/11/ 51:35-13 c6m B;lake-Baham Baasi nN, ort hAtlant iOc cean. FIGURE 8S9,.—Zon Pe aD, SD PSit 5e77/12/5 1:15-11 c7m S;hatsk Ryis en,orthwester nPacif iOc cean. NUMBE 8R5 233 234 SMITHSONIA CNONTRIBUTION TS POALEOBIOLOGY PLATE 64 Globoconus adaubjergensi s(Bronnimann 1,953) (Figure s1-7 ,10-12 :bar s= 50 pm ;Figure s8 ,9 :bar s= 10 pm) FIGURE S1-3 5 ,6 ,8 ,9, .—Zon ePic B, rightsea Ftm .M, aryland F; igur e 8(view o uf ltimat echambe ro Ff igur e6) an dFigur (9evie wo ufltimat cehambe o rFfigur 5e s)howin gmicroperforat we a tllextur aen dporeles sharp pustules. FIGUR 4E.—Zon Pe 0B,razo Rsive Tr,exas. FIGUR 7E.—Zon Pe aM, iller Fserr yA,labam ac,or 2e2 5s,amp l2e56. FIGURE 1S0-1 2—. Zon Pei cM, idwa Fym T.,exa sP,lumme srtatio n14. NUMBE 8R5 235 236 SMITHSONIA CNONTRIBUTION TS POALEOBIOLOGY PLATE 65 Parvularugoglobigerina alabamensis (Liu and Olsson ,1992) (Figure s1-5 :bar s= 50 pm ;Figure 6 :ba r= 10 jam) FIGUR E1.—Zon eP l Ma, iller Fserry A, labama c,or e225 s,ampl e265. FIGURE S2 3,.—Zon ePla M, iller Fserr yA, labama c,or e225 s,ampl e266. FIGUR 4E.—Zon Pel aM, iller Fserr yA,labam ac,or e22 6s,amp le20. FIGURE S5 6,.—Zon ePla M, iller Fserr yA, labama F;igur e5 h,olotype s,ampl e4 0fee atbov ePrairi eBlu fCfhalk; Figur 6ev,ie wo wf a o l3lfr dchambe o rhfolotyp sehowin gmicroperforat peor me oun dtexture. Parvularugoglobigerina extensa (Blow 1, 979) (bar s=5 0pm) FIGURE 7S9,-12.—Zon Pe aD, SD PSit 5e77/12/5 9:4-9 c6m S;hatsk Ryis en,orthwester nPacif iOc cean. FIGUR E8 —. Zon ePa E ,Kl e fT,unisia. FIGUR 1E3.—Zon Pe aty, p leev eo Plef.ugubin aG,ubb isoectio nc,entr aAlppennine sIt,aly. NUMBE 8R5 237 238 SMITHSONIA CNONTRIBUTION TS POALEOBIOLOGY PLATE 66 Parvularugoglobigerina eugubina (Luterbacher and Premol iSilva ,1964) (Figure s1-4 6, -12 :bar s= 50 |im ;Figure 5 :ba r= 10 |im) FIGURE S1-5.—Zon ePa M, iller sFerry A, labama c,or e225 s,ampl e328 F;igur e5 v,iew o wf a lol 2fn dchamber o Ffigur s2ehowin mg icroperforat we a tlel xtur we it fahe wincipien ptor me ounds. FIGURE S6 8 ,9 ,1, 2.—Zon ePa D, SD PSit e577/12/5 1: 15-11 7cm. FIGURE S7 1,0.—Zon ePa E ,Kle fT,unisia. FIGUR 1E1.—Zon Pe aD,SD SPit 5e77/12/ 51:34-13 c6m S;hatsk Ryis en,orthwester Pnacif iOc cean. NUMBE 8R5 239 240 SMITHSONIA CNONTRIBUTION TS POALEOBIOLOGY PLATE 67 Parvularugoglobigerina eugubina (Luterbacher and Premol iSilva ,1964) (Figure s1-10 1, 2 1, 3 :bar s= 50 (am ;Figure s11 1, 4 :bar s= 10 pm) FIGURE S1-9 —. Zon ePa t,yp eleve o lP fe.ugubina G, ubbi osection c,entra Alppennine sI,taly. FIGURE S10-12.—Zon ePa D, SD PSit e577/12/5 1:25-12 6cm S;hatsk yRise n,orthwester nPacif iOc cean; Figur 1e 1v,ie wo Ffigur 1e s0howin rgecrystallize wda tlhl aotbscure ms icroperfora twea tlel xture. FIGURE S13 1,4.—Zon ePa M, iller sFerry A, labama c,or e226 s,ampl e20 F;igur e14 v,iew o wf a lol 4ft hchamber oFfigu r1 es3howin mg icroperfora twea tlel xtu raen sdcattere pdo rme ounds. NUMBE 8R5 241 mm SB ,,C,-a 242 SMITHSONIA CNONTRIBUTION TS POALEOBIOLOGY PLATE 68 Woodringina claytonensis Loeblich and Tappan ,1957 (Figure s1-5 :bar s= 50 (am ;Figure 6 :ba r= 10 pm) FIGUR 1E.—Zon Pe 0B,razo Rsive Tr,exas. FIGURE S2 6,.—Zon ePla D, SD PHol e390A/11/5 1:35-13 6cm B; lake-Baham aBasin N, ort hAtlant iOc cean; Figur 6ev,ie wo 2fn dchambe o rFfigur s2ehowin gmicroperforat we a tllextur aen dblun ptustules. FIGUR E3 —. Zon ePa M, iller Fserr yA, labama c,or e226 s,ampl e18. FIGUR 4E.—Zon Pel aM, idwa Gyrou pT,exa sP,lumme srtatio 4n. FIGUR 5E.—Zon Pe aD,SD SPit 5e77/12/ 51:15-11 c7m S;hatsk Ryis en,orthwester nPacif iOc cea ns,pecimen show iingtermedi amteorpholo wg Giytuhembeli tcrriaetacea. Woodringin ahornerstownens iOs lsson 1,960 (bar s=5 0pm) FIGURE 78S,.—Topotype Zs,on Pe 3H,ornerstow Fnm N.,e wJersey. FIGURE S9 1,0 1,3 1,4.—Zon ePa D, SD PSit e577/12/5 1:13-11 4cm S;hatsk Ryise n,orthwester nPacif iOc cean. FIGURE 1S 11,2.—Zon Pei cM, idwa Fym T.,exa sP,lumme srtatio n14. NUMBE 8R5 243 vHtttf&PiW D’Hondt, Steven, Liu, Chengjie, and Olsson, Richard K. 1999. "Family Guembelitriidae Montanaro Gallitelli, 1957." Atlas of Paleocene planktonic foraminifera 85, 77–88. View This Item Online: https://www.biodiversitylibrary.org/item/267464 Permalink: https://www.biodiversitylibrary.org/partpdf/352117 Holding Institution Smithsonian Libraries Sponsored by Smithsonian Institution Copyright & Reuse Copyright Status: In copyright. Digitized with the permission of the rights holder. Rights Holder: Smithsonian Institution License: http://creativecommons.org/licenses/by-nc-sa/4.0/ Rights: http://biodiversitylibrary.org/permissions This document was created from content at the Biodiversity Heritage Library, the world's largest open access digital library for biodiversity literature and archives. Visit BHL at https://www.biodiversitylibrary.org. This file was generated 27 May 2023 at 22:39 UTC