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Gradstein, F. M., Ludden, J. N., et al., 1992 Proceedings of the Ocean Drilling Program, Scientific Results, Vol. 123

16. LOWER NANNOFOSSIL OFF NORTHWESTERN AUSTRALIA (LEG 123)1

Jörg Mutterlose2,3

ABSTRACT

Moderately to sparsely nannofossiliferous Neocomian siliciclastics and rich - nannofossil chalks were cored at two Leg 123 sites on the abyssal plains off northwestern Australia. At Site 765, the basal 70 m of cored section yields questionable and to early assemblages of moderate diversity containing Cruel- ellipsis cuvillieri, Tegumentum striatum, Speetonia colligata, and Crucibiscutum salebrosum. The overlying Hauterivian- lower Aptian is represented by 140 m of sediments barren of nannofossils. Above this, the remaining 80 m of the Lower Cretaceous section has been assigned to the Rhagodiscus angustus Zone (late Aptian-early Albian in age) and the Prediscosphaera columnata Zone (middle-late Albian in age). Common species include Rhagodiscus angustus, Predis- cosphaera columnata, Eprolithus floralis, Eprolithus sp., Chiastozygus litterarius, Rucinolithus irregularis, and Flabel- lites biforaminis. At Site 766, the Neocomian, represented by 200 m of sediment, yields C. cuvillieri, T. striatum, S. colligata, and C. salebrosum. Within the overlying Aptian-Albian sequence of 80 m, the Rhagodiscus angustus, and P. columnata zones were recognized. The paleobiogeographic patterns and implications are discussed, with special emphasis paid to the bipolar high-latitude distribution pattern of C. salebrosum in the -Hauterivian. Biostratigraphically important species are discussed and their occurrence in the Indian Ocean is compared with one from the Tethys and Boreal realms. Two new species, Serbiscutum gaultensis and Eprolithus bettenstaedtii, are described.

INTRODUCTION At both sites, scientists recovered thick and apparently com- plete Lower Cretaceous sequences. Only in the southern oceans During Legs 122 and 123, eight holes were drilled (Sites 759 (at Site 261) had a fairly consistent sequence of Neocomian through 766) off northwestern Australia across the Exmouth Pla- sediments previously been drilled. Sediments of an Aptian-Al- teau and the adjacent abyssal plains. These sites cover two com- bian age, however, have been recovered in several ODP/DSDP plete transects from a continental margin to an old ocean basin sites (Fig. 3). Thus, Sites 765 and 766 offer scientists an ideal (Fig. 1). opportunity to gain firsthand information about the Neocomian One of the principal goals of Leg 123 (Sites 765 and 766) was faunas and floras of the southern oceans. In reference to calcare- to decipher the early rifting history of the northwestern Australian ous nannofossils, the following aspects are of particular interest: margin and the initial opening of the Indian Ocean by drilling the oldest sediments and basement of the Indian Ocean in the Argo 1. Biostratigraphy of the Neocomian in the southern oceans. Abyssal Plain. 2. The Aptian floral turnover, which has been observed else- Drilling at Site 765, which is situated at the southern margin where. of the Argo Abyssal Plain, recovered a thick (330 m) sequence of 3. Paleobiogeography of Neocomian floras. Lower Cretaceous and possibly Uppermost sediments, 4. Comparison of Tethyan, Boreal, and Indo-Pacific floras. covered by 600 m of Upper Cretaceous and Tertiary sediments. The Lower Cretaceous sediments are dominated by siliciclastics, METHODS in particular, varicolored claystone (lithologic Units VII, VI, V, Simple smear-slide preparations were examined under the and Subunit IVD) in the lower part ([?]Tithonian, Berriasian- light microscope using a magnification of 1500x. The abundance early Aptian) and nannofossil chalks (late Aptian-Albian) in the of calcareous nannofossils in the Lower Cretaceous material is upper part (lithologic Subunits IVC through IVA; Fig. 2). variable. Nannofossils may constitute anywhere from 0% to 70% Site 766, farther south at the boundary between the Gascoyne of the rock. Abundances for each species were estimated by using and Cuvier abyssal plains, penetrated about 300 m of Lower the following criteria: Cretaceous sediments (Valanginian-Albian). These consist of sandstones and siltstones in the lower part (Valanginian-Barre- VA = very abundant (11-100 specimens/field) mian) and nannofossil chalks in the upper part (Aptian-Albian; A = abundant (2-10 specimens/field) Fig. 2). The Lower Cretaceous sediments are overlain by 160 m C = common (1 specimen/1-10 fields) of nannofossil oozes of , Tertiary, and Quaternary F =few (1 specimen/11-100 fields) age. R = rare (1 specimen/more than 100 fields)

Preservation is indicated in the following way: El (slightly etched), E2 (moderately etched), and E3 (heavily etched). Al- 1 Gradstein, R.M., Ludden, J. N., et al., 1992. Proc. ODP, Sci. Results, 123: though it occurs, overgrowth was not noted in the range charts. College Station, TX (Ocean Drilling Program). For examination under the scanning electron microscope, samples 2 Department of Geology, University of Nebraska, Lincoln, NE 68588, U.S.A. Current address: Institut fur Geologie und Palàontologie, Universitàt Han- were prepared following the procedures outlined by Wise and nover, Callinstrasse 30, 3000 Hannover 1, F.R.G. Kelts (1972). J. MUTTERLOSE

\ Scott 12°S Plateau

Argo Abyssal Plain m765S h>260 Gascoyne\r59-761, 7641/Swan Canyon M^jf /Aöyssa/ Plain 766-^ ^::-; WESTERN AUSTRALIA

• ODP Leg 123 Sites A ODP Leg 122 Sites 24° o DSDP Sites

Figure 1. Location of Sites 765 and 766 in relation to DSDP Leg 27 and ODP Leg 123 sites.

Many of the Neocomian siliciclastics recovered from Sites 765 was penetrated, of which about 330 m has been assigned an Early and 766 are barren of nannofossils. Although much of this mate- Cretaceous (Berriasian-Albian) and possibly rial has been examined, barren samples only occasionally are ([?]Tithonian) age for the basal few meters. Abundances and reported in the range chart. Thus, not all samples considered here preservation of nannofossils are shown in Table 1. are listed in the range charts. Basaltic basement was encountered at a depth of 931.2 m in Taxa considered here are listed in the Appendix. Most biblio- Section 123-765C-62R-4. The lowermost part of the overlying graphic references for these taxa are given in Perch-Nielsen sedimentary sequence, Sections 123-765C-62-4 to 123-765C- (1985); any not included therein are given in the references. All 58-2 (lithologic Subunits VIIB and VIIA), apart from a few illustrations in the plates are scanning electron photomicrographs samples, are barren of calcareous nannofossils. Samples taken unless noted otherwise in the captions. In the captions, P, D, and directly above the basalt (123-765C-62R-4, 28 cm; -62R-4, 10 cm; L denote proximal, distal, and lateral views, respectively. Abbre- -62R-3, 100 cm; -62R-3, 100 cm; -62R-3, 20 cm; -62R-2, 140; viations LM, TL, POL, and PH denote light photomicrograph, -62R-2, 28 cm) are barren. The first nannofossils were observed plain transmitted light, cross-polarized light, and phase contrast in Sample 123-765C-62R-1, 120 cm. Samples 123-765C-61R-4, light photomicrograph, respectively. 30 cm, and 123-765C-61R-3, 52 cm, yield similar, poorly pre- LOWER CRETACEOUS NANNOFOSSIL served, low-abundance, low-diversity assemblages. These impov- erished floras are dominated by Cyclαgelosphαerα deflαndrei BIOSTRATIGRAPHY with less common Microstαurus chiαstius and Microstαurus All the Lower Cretaceous zonation schemes published so far quαdrαtus. These stratigraphically older specimens of C. deflαn- are of limited value, because they refer to the Tethys (Sissingh, drei are large and oval, while younger forms in the overlying 1977; Thierstein, 1971, 1973; Roth, 1978; Covington and Wise, Neocomian section are circular. None of these species can be used 1987; Bralower et al., 1989; Applegate et al., 1989) or the Boreal to define the age of these samples exactly. Realm (Jakubowski, 1987; Mutterlose, 1988; Crux, 1989; Mutter- Based on sections from southern Europe and the southern lose, 1991). A zonation based on material from the southern North Atlantic (Bralower et al., 1989), M. quαdrαtus has its first oceans has been published only for the Aptian-Albian (e.g., Wise, occurrence at the base of the Tithonian, whereas M. chiαstius is 1983). The zonation presented in this chapter, however, does use not present until the middle Tithonian. Both species extend into age determinations from a combination of the published schemes. the Hauterivian and , respectively. C. deflαndrei has A detailed discussion of the zonation scheme used here is pre- been used by Roth (1983) as an index fossil that ranges from the sented later. middle to the Valanginian. Thus, none of the species observed in these lowermost nanno- Site 765 (Table 1) fossil-bearing samples gives any decisive clue for the exact age Hole 765C (15°541'S, 117°34.495'E; water depth, 5717 m) is of the sediments encountered above the basement. However, the located about 350 km northwest of the coast of western Australia exact age of these sediments is a crucial question, because a in the southernmost part of the Argo Abyssal Plain, about 50 km to age had been expected before drill- north of the (Fig. 1). About 931 m of sediment ing. One of the objectives of this site was to encounter the oldest

344 LOWER CRETACEOUS NANNOFOSSIL BIOSTRATIGRAPHY Site 766 Site 765 ID CD Naπno- O Nanno- 3 plankton 3 O plankton cr cr o CD Age Lithology Events Age Lithology Events

650- \-^T \ ooze, 9 Chalk P. P. 200 columnata columnata silt II— 1I ston' e 700-

R. angustus R. yj basalt 250- E. floralis angustus C. achylosus J FAD F. biforaminis 750 π LAD C. cuvillieri 300- - S. colligata α> 800- I

350-

850- el early Haut 7. striatum 400- o late Valaπg Valanginiaπ j T. striatum \\ 900- C. cuvillieri T. striatum V. matalosa

450 T. striatum C. cuvillieri 7^--s rAA C. deflandrei V. matalosa Figure 2. Lithologic and biostratigraphic summary of the Lower Cretaceous of Holes 765C and 766A.

oceanic crust in the Indian Ocean. A pre-drilling assumption of a Site S. Atlantic S. Ocean Indian Ocean Kimmeridgian-Oxfordian age for the oldest sediments overlying

O T- CO C\J the basement was based on two observations: CM co •>- σ> σ> CO CD CD CD CD CD CO CO f CD CD c\i r-^ t~^ i~~ r^ r~. Albian 1. Paleomagnetism indicated a Late Jurassic age for the base- ment (anomaly M26). Aptian η 2. At Hole 261 (located about 320 km farther north and thus Barremiaπ having a presumably younger basement), sediments of Oxfordian and Kimmeridgian age were encountered above the basement. Hauterivian This implied an even older age for the crust at Site 765 farther Valaπginiaπ south. Berriasian According to these assumptions, the initial opening of the Tithonian Indian Ocean should have happened during the Oxfordian or Callovian. Because no other index species to indicate a definite Kimmeridgian Tithonian or Berriasian age have been observed in Hole 765C, the Oxfordian age of the oldest sediments is questionable. Support for a Titho- Callovian nian age of the sediments in Sections 123-765C-62R-1 through -61R-3 comes from correlation of the nannofossil assemblages Figure 3. Location and ranges of DSDP/ODP drill sites in the South consisting of C. deflandrei, M. chiastius, and M. quadratus, with Atlantic, Southern, and Indian oceans that penetrated Lower Cretaceous a similar assemblage in DSDP Site 261, 320 km north of Site 765. sediments. In Site 261, this assemblage occurs above Kimmeridgian to lower

345 * Table 1. Distribution of Lower Cretaceous calcareous nannofossils, Hole 765C.

Nanno- plankton 1 Cyclagelosphaera deflandrei 5 Microstaurius quadratus 8 Zeugrhabdotus embergeri 3 Cyclagelosphaera sp . 6 Cyclagelosphaera margerelii 9 Vagalapilla matalosa 2 Watznaueria barnesae 4 Microstaurius chiastius 1 Ellipsagelosphaera fossacincta 1 Crucibiscutum salebrosum 1 2 Cruciellipsis cuvillieri 1 3 Ethmorhabdus hauterivianus 1 4 Haquis circumradiatus 1 5 Lithraphidites carniolensis 1 6 Manivitella pemmatoidea 1 7 Rucinolithus wisei 1 8 Tegumentum sp . 1 9 Vekshinella cf . stradneri 1 0 Biscutum constans 2 0 Watznaueria ovata 2 1 Zeugrhabdotus erectus ssp . 2 9 Axoporhabdus dietzmannii 3 1 Cretarhabdus sp . 3 2 Grantarhabdus meddii 3 Micrantholithus (frags. ) 3 6 Rotelapillus laffittei 3 7 Tegumentum octiformis 3 8 Glaukolithus diplogrammus 2 Conusphaera rothii 2 3 Hemipodorabdus gorkae 2 4 Tegumentum striatum 2 6 Tubodiscus sp . 2 7 Watznaueria britannica 2 8 Assipetra infracretacea 3 0 Cretarhabdus angustiforatus 3 4 Reinhardtites fenestratus 3 5 Rhagodiscus asper 3 9 Chiastozygus tenuis 4 0 Vekshinella stradneri ssp . 2 4 1 Cretarhabdus 3 element s Age zone Preservatio n Sample no. 0 5 E2 31, CC . A . R . . . . R C R F F . F . . . . ft. ft. 0 5 . C . F R E2 32-1,27 . A . F . R . F R F F R R . C . . R . . R E2 32-2, 53 . C . . . . R R R . . R F R R R E2 32-3, 53 . F . E3 32, CC . R E2 33-1,61 . A . R . R C C . . R . . . F R R R R early-late P. E2 33-3, 30 . F . . . . F R R . R R R Albian columnata E2 33, CC . R ft. 0 5 O 5 0 S Qi • Qi El 34-2, 70 . A . R . R A F R F . . . R C oi ft. a . . . C R . R . . R El 34-4, 15 . C . R . R C F R F . . . R F . . . C R . R . . R

E2 34, CC . C . R . . R R R F . R 70 7 0 T i R R R . R . R R El 35-1,50-51 . VA . R . R C R R C . . . R C R . . R R R . F . . R R R . . . . R C R . . . . R E2 35-1, 105 . A . R . C F . C . C R . F . . F F R . R . . . E2 35-2, 47-48 . C . R . . F R R R F R R R . R R . . R F R . R . . R E2 35-4, 65-66 . A . R . R A R R . . R R R . R R F E3 35, CC . C . . . C R R R R R R E2 36-1, 136 . VA . R . R A R R F . F . R . R . F R F R . R E2 36-5, 116 . C . R . R A F R . . . . R F . . . . R . . R . . R . . R R . . . F . E2 36-6,51-53 . A . R . . C R R F R . . . R . R . R . R R . F R . R . R . E2 37-2, 102 . VA . R C C F F . R F . R C R . R . . . β 5 •

El 37-4,21-22 .A ... F C R F C R . . . R • 0 5 • ft. a ; o s F . R R . . . . R R . . . R R E2 38-1,20 . VA . . . R A F R C F . . . F R . . R . . . . C R . F . . R

late Aptian- E2 38-2, 20 . A . R . R A R R F . . . . R 70 • 70 J O R R R R. angustus early Albian E2 38-3, 135 . A . R . F C R R F R . . R F R F F R . F . R . E2 38-6, 45 . A . R . . A R R C R . . R R . R . R R R R . R . R . E2 39-1, 140 . VA . R . R A F R C . R R R . . R F R R F R . R . R . El 39-2, 110 . VA . R . . A R R C . F R . . R R . R . . . . F R . F . . R El 39-4,30-31 . A . R . C A R R C . . . R F R . . R F . . F . . . R . C R . R . . R 39-5, 4-5 El . VA . F . R A F R F . . . R C R . . . R . R . - d j o 70 • R . F R . F . R F El 40-1,20 . A . R . . A R R F R R R C R . R R R R E2 40-2, 106-107 . A . R . . A R R R . R . R . R R . R . R F R . R . . R E2 40-3, 002 . A ... R A . F F . R . . . R F . R . . . R . . . R F R . R R . R E2 40-4, 30 . A . R . . C C . . . R R R E2 40-4, 128 . F . . . E2 40-5, 47-48 F 41-1,49-50 42-1,60-61 . R E3 Hauterivian- 42-2, 50 Aptian 42-3, 7-9 42-4,51-52 . R . . . . R . E2 42-5, 45-47 43-6, 69 44-1,23-24 . F ...... R R . . . . E2 Table 1 (continued).

E2 44-3, 149-150 . R . . . . R 44-4, 14_15 45-1,95 45-2,61-62 45-2, 65 48-3, 30 Hauterivian- 48-4, 50 Aptian 52-1,95 E3 52-2, 66-69 . F . . . . R . R . . . . 52-3, 65 54-3, 85 E3 54-4, 57_58 F R R . R E2 55-1, 120 . A ... F C F R . . . . R F . R R . R . E2 55-2, 109-110 R A . R . R A C C . F R R . F R . . . R . R R El 55-3, 50 . A ... F C F C R C . . R F R R R R R late E3 55-4,61-62 . C . . . R F R F . . R R . R R . Valanginian- E2 56-1,85 . C . . . R C F R R F . R early E2 56-2, 60-61 . VA . R . F A C C R F F . R C R . . . F . R R . R . . . . . R Hauterivian E2 56-3, 137 R A . R . F C F F R R R . . F . R . R . . F . . . . R . E2 56-4, 10 R A . R . C A C F R F R . R F . . . R R R R . R . . F . R . E2 57-1, 135 . C . . . C A C . . . R . R F . R R . R T. striatum E2 57-2, 109-111 . VA . . . C A F R R R R . . F . R R E2 57-3, 110 R A . . . F A R R . C R . R F . R . . R R . . R . . R . . R R . . R R R . E2 57-3, 140-150 R A . . . C A F C . A R . R F R R . . R . R . . . R R R R R R R R F R . E2 57-4, 78-79 . A ... R A C . R . F . F . . R . R . . . Valanginian E2 57-5, 14-16 F E2 57-6, 30 R E2 57-7, 30 F E2 58-1,85 R VA . R . R A C . R F F . R F R R . F F . R R F R F ... E2 58-1, 131 R A . R . F C F . R F R R R F R F R R R F . . . . E3 58-2, 140 F R E2 58-3, 140 . F . . . R . . . E2 58-4, 120-122 . F . . . . R . R . E2 59-3, 82-84 . F . . . R R R . . Berrias.- 59-4, 90 Valan. 60-2, 50 E3 60-3, 90 . R . E3 60-4, 100 . R . 60-5, 85-87 61-1,30 61-2,30 E2 61-3,52 F . R E2 61-4, 30 C . F F F 61-4, 107 E2 62-1,65 A .... E3 62-1, 120 C R . . . (?) Tithonian 62-4, 10 62-4, 28

Note: Abundances are characterized by VA (very abundant) = 11-100 specimens/field, A (abundant) = 2-10 specimens/field, C (common) = 1 specimen/1-10 fields, F (few) = 1 specimen/11-100 fields, R (rare) = 1 specimen/more than 100 fields. Preservation: El = slightly etched, E2 = moderately etched, and E3 = heavily etched. Table 1 (continued).

| Nanno- u plankton g Age zone o- Sample no. E2 31, CC E2 32-1,27 E2 32-2,53 E2 32-3,53 E3 32, CC E2 33-1,61 early-late P. E2 33.3,30 Albian columnata ^j 33 cr El 34-2,70 El 34-4, 15 E2 34, CC El 35-1,50-51 E2 35-1, 105 E2 35-2,47-48 E2 35-4,65-66 E3 35, CC E2 36-1, 136 E2 36-5,116 E2 36-6,51-53 E2 37-2, 102 El 37-4,21-22 E2 38-1,20 lateAptian- „ E2 38-2,20 early Albian Ran8ustus E2 38-3,135 E2 38-6,45 E2 39-1, 140 El 39-2,110 El 39-4,30-31 El 39-5,4-5 El 40-1,20 E2 40-2, 106-107 E2 40-3,002 E2 40-4,30 E2 40-4, 128 E2 40-5,47-48 41-1,49-50 E3 42-1,60-61 Hauterivian- 42-2,50 Aptian 42-3,7-9 E2 42-4,51-52 42-5, 45^7 43-6, 69 E2 44-1,23-24 Table 1 (continued).

E2 44.3, 149-150 44-4, 14_15 45-1,95 45-2, 61-62 45-2, 65 48-3, 30 Hauterivian- 48-4, 50 Aptian 52-1,95 E3 52-2, 66-69 52-3, 65 54-3, 85 E3 54_4; 57_58 E2 55-1, 120 R . . . E2 55-2, 109-110 R R El 55-3, 50 R . R late E3 55-4, 61-62 Valanginian- E2 56-1,85 early E2 56-2, 60-61 . R R Hauterivian E2 56-3, 137 E2 56-4, 10 . F R R . E2 57-1, 135 T. striatum E2 57-2, 109-111 R . . . . E2 57-3, 110 E2 57-3, 140-150 E2 57-4, 78-79 Valanginian E2 57-5, 14-16 E2 57-6, 30 E2 57-7, 30 E2 58-1,85 E2 58-1, 131 E3 58-2, 140 E2 58-3, 140 E2 58-4, 120-122 E2 59-3, 82-84 59-4, 90 Berrias.- 60-2, 50 Valan. E3 60-3, 90 E3 60-4, 100 60-5, 85-87 61-1,30 61-2,30 E2 61-3,52 E2 61-4,30 61-4, 107 62-1,65 (?) Tithonian E2 E3 62-1, 120 62-4, 10 62-4, 28 J. MUTTERLOSE

Tithonian sediments and below Berriasian to Hauterivian sedi- co-occurrence of the two species may result from reworking as ments (Proto Decima, 1974). In view of this correlation and well. because of the absence of species typical for the Berriasian, I have Based on the highest occurrence of R. wisei in Sample 123- assigned a Tithonian age to the lowermost nannofossil-bearing 765C-55R-2, 109-110 cm, Sections 123-765C-56R-4 to -55R-2 samples of Site 765 from Sections 123-765C-62R-2 to -61R-3. (lithologic Unit VI) are early-late Hauterivian or older in age. The subsequent interval, including Samples 123-765C-61R-2, This is further constrained by the occurrence of Speetonia 30 cm, to -58R-2, 140 cm, is difficult to date using calcareous colligata in Sample 123-765C-56R-2, 60-61 cm. In other areas, nannofossils, because definite index species are missing. A Titho- S. colligata ranges from the base of the Berriasian to the early-late nian(?) and/or Berriasian-Valanginian age is possible, based on Hauterivian (Bralower et al., 1989; Mutterlose, 1991). samples from the underlying and overlying sections. A Berriasian The acme of C. salebrosum, observed in Samples 123-765C- age is constrained by dinoflagellate markers, which suggests a late 58R-1, 131 cm, to -55R-2, 109-110 cm, may possibly be another Berriasian age {Batioladinium reticulatum Zone) for Samples age indicator. In the Boreal Realm, C. salebrosum is most com- 123-765C-59-5, 8-9 cm, and -59R-4, 105-112 cm (McMinn, this mon in the late Valanginian and early Hauterivian, while it is rare volume). only in the late Hauterivian and . These observations Sample 123-765C-58R-1, 131 cm, yields the first diverse are supported by the highest occurrence of T. striatum in Sample nannofossil assemblage, which is dominated by Watznaueri 123-765C-55R-3, 50 cm. barnesae. Above this, Samples 123-765C-55R-1,120 cm, to -40R-4, 128 Biostratigraphically important species include Rucinolithus cm (lithologic Units VI and V) contain only rare W. barnesae and wisei, Crucibiscutum salebrosum, Cruciellipsis cuvillieri, Lithra- a few other species. This interval was assigned a late Hauterivian phides carniolensis, and Vagalapilla matalosa. to early Aptian age; this age determination is based, however, on The biostratigraphic information supplied by the most impor- data only from the underlying and overlying beds. This assump- tant marker, R. wisei, is biased by the different ranges given by tion is constrained by palynological data (McMinn, this volume). Thierstein (1971; Berriasian-Valanginian) and Bralower et al. Samples 123-765C-54R-CC to -54R-4, 49-50 cm, contain dino- (1989; upper Berriasian-lower Valanginian). Based on more re- flagellates of a late Hauterivian to Barremian age. The highest cent publication, a late Berriasian to late-early Valanginian age is occurrence of palynomorphs characterizing the late Barremian assumed for R. wisei. The oldest occurrence of C. salebrosum has was observed in Sample 123-765C-50R-CC. The interval 123- been reported from the Peregrinocceras albidum ammonite Zone 765C-49R-CC to 40R-4 has been assigned an early Aptian age on (late Ryazanian) from Speeton, where this species is common the basis of palynomorphs. (Crux, 1989). The Ryazanian corresponds to the upper part of the A well-preserved nannoflora occurs in Sample 123-765C- Berriasian (for detailed discussion of this problem, see Zeiss, 40R-4, 30 cm. A fairly rich and diverse assemblage consisting of 1983). However, because no older nannofossil-bearing strata Rhagodiscus angustus, Chiastozygus litterarius, Eprolithus flo- were examined, C. salebrosum may extend into the lower Berri- ralis, (= Eprolithus apertior) sp., and Flabellites biforaminis asian. suggests a late Aptian to earliest Albian age. The first occurrence Thus, based on the co-occurrence of this species with R. wisei, of R. angustus has been used widely to define the base of the a late Berriasian to late-early Valanginian age of this sample Rhagodiscus angustus Zone (sensu Thierstein, 1971), which has seems most likely. C. cuvillieri ranges from the latest Tithonian a late-early to earliest Albian age (Perch-Nielsen, 1979, 1985; to the early-late Hauterivian (Bralower et al., 1989) in the Medi- Jakubowski, 1987). The other species mentioned have their first terranean area. The species is rare only in the Boreal Realm, where occurrences in the early Aptian (Mutterlose, 1991). Thus, the it ranges into the late-early Hauterivian. Abundances and preser- Chiastozygus litterarius Zone (sensu Thierstein, 1973) Rucino- vation of nannofossils are shown in Table 2. A sequence of lithus irregular is Zone, sensu Thierstein, 1977) of early Aptian interlayered C. cuvillieri ranges from the latest Tithonian to the age is represented by an underlying interval barren of nanno- early-late Hauterivian (Bralower et al., 1989) in the Mediter- fossils or is missing. ranean area. This species is rare only in the Boreal Realm, where Sample 123-765C-35R-1, 105 cm, is characterized by the it ranges into the late-early Hauterivian. L. carniolensis has its lowest occurrence of small-sized Prediscosphaera columnata, first occurrence in the uppermost Tithonian, near the Titho- which first appear in the early Albian elsewhere. nian/Berriasian boundary (Bralower et al., 1989). The presence of V. matalosa is of interest because this species has so far been Site 766 (Table 2) recorded only from the latest Barremian onward. Hole 766A (19°55.985'S, 110°27.243'E; water depth = 4008 Sample 123-765C-58R-1, 85 cm, yields an assemblage com- m) was drilled at the boundary between the Cuvier and Gascoyne parable to that of Sample 123-765C-58R-1, 131 cm. However, abyssal plains about 350 km off the coast of western Australia Tegumentum striatum (= Eiffellithus striatum) is of stratigraphic (Fig. 1). This site is located about 650 km southwest of Site 765. importance. For the Mediterranean area (Sites 603, 638), Apple- About 460 m of sediment was encountered, of which about 300 gate et al. (1989) reported the first occurrence of this species from m (Cores 123-766A-48R to -18R) has been assigned an Early the earliest Hauterivian. For the Boreal Realm, Mutterlose (1991) Cretaceous age (latest Valanginian to Albian). Abundances and already described T. striatum from the early-late Valanginian; preservation of nannofossils are shown in Table 2. A sequence of however, as a result of facies control, it is absent from the early interlayered, small basalt sills and sediments was encountered at Valanginian. Thus, this species may extend into the early Valan- a depth of 458 mbsf (Core 123-766A-48R). This has been replaced ginian. In the Boreal Realm, T. striatum extends into the late by a thick diabase intrusion at 467 m (Core 123-766A-49R). Hauterivian; however, it becomes rare in the late Hauterivian. The lowermost nannofossil-bearing Sample 123-766A-50R-1, Based on the co-occurrence of T. striatum and R. wisei, this 5 cm, yields a fairly rich and diverse assemblage dominated by sample may have a late-early to early-late Valanginian age. W. barnesae. Biostratigraphically important species include C. Based on the highest occurrence of R. wisei in Sample 123-765C- cuvillieri, C. salebrosum, and Tegumentum. The presence of the 57R-1, 135 cm, the interval from -58R-1 to -57R-1 has been latter species and the absence of R. wisei, observed in Hole 765C, assigned a Valanginian age. The calcareous intervals from which indicates a late Valanginian age. While C. cuvillieri and C. sale- R. wisei has been recorded have been interpreted as classical brosum have their first occurrences in the Berriasian (for discus- turbidite sequences (Ludden, Gradstein, et al., 1990). Thus, the sion, compare with Site 765), T. striatum is known from the early

350 LOWER CRETACEOUS NANNOFOSSIL BIOSTRATIGRAPHY to late Valanginian onward. On the basis of the last occurrence of previous discussion). These show similar impoverished floras that T. striatum in Sample 123-766A-45R-5, 97-99 cm, interval 123- consist of C. deflandrei, M. chiastius, andM. quadratus only. The 766A-50R-1 to -45R-5 was assigned a late Valanginian to early Berriasian- Hauterivian interval of Site 261 yielded C. cuvillieri Hauterivian age. and C. salebrosum. The presence of these species allows for a Based on the highest occurrence of C. cuvillieri in Sample correlation of Sections 27-261 -31R-2 to -28R-2,123-765C-58R-1 123-766A-31R-3, 29-31 cm, Sections 123-766A-49R-3 to -31R-3 to -55R-2, and 123-766A-50R-1 to -31R-3. are early-late Hauterivian or older in age. C. cuvillieri has an PALEOENVIRONMENT AND earliest to late Hauterivian extinction elsewhere. PALEOBIOGEOGRAPHY Samples 123-766A-31R-1, 39-41 cm, to 123-766A-27R-3, 18-19 cm, have been assigned a latest Hauterivian and Barremian Lower Cretaceous nannofossils show provincialism similar to age. However, this age assignment is based only on the presence that exhibited by macrofaunas (ammonites, belemnites, bivalves, of C. cuvillieri in the underlying sections and of typical Aptian , and foraminifers). Major faunal and floral changes species in the overlying sections. The base of the Aptian is defined across the Barremian/Aptian boundary caused a reorganization of by the lowest occurrence of Corollithion achylosus, Flabellites the paleogeographic distribution patterns separating the Berria- biforaminis, Rotelapillus crenulatus and in E. floralis in Sample sian to Barremian (= Neocomian) from the Aptian to Albian (= 123-766A-27R-1, 108-109 cm. Interval 123-766A-27-1 to -26-2 ). These changes, referred to as the worldwide Aptian turn- is of early Aptian age. The lowest occurrence of R. angustus over, revitalize the outdated terms "Neocomian" and "Gault" used (indicating a late-early to early-late Aptian age) is found in here. For the Berriasian to Barremian interval, it is possible to Sample 123-766A-26R-1, 10 cm. The R. angustus Zone (late differentiate two realms as follows: Aptian-earliest Albian) is represented by the interval 123-766A- 26R-1 to -22-CC. The nominate species for the overlying P. 1. A northern, moderate temperate Boreal Realm (north- columnata Zone (early-late Albian) first occurs in Sample 123- western Europe, Russia, Greenland, northern America): Charac- 766A-21R-CC. terized by cosmopolitan, Boreal, endemic, and scattered incur- sions of Tethyan floras. Endemic species are those forms that Regional Correlation evolve from originally Tethyan species, but become restricted to Because drilling at Sites 765 and 766 recovered a fairly com- the Boreal Realm (e.g., Nannoconus abundans, Nannoconus bo- plete Lower Cretaceous sequence moderately rich in calcareous realis). The scattered incursions of Tethyan floras mark high- nannofossils, a biostratigraphic correlation between both sites stands of sea level and warming periods. (based on calcareous nannofossils) is possible (Fig. 2). Because 2. A southern, tropical-to-subtropical Tethyan Realm of the restricted environment, most of the Neocomian sediments (southern and central Europe, the Mediterranean area, Mexico): consist of siliciclastics that were deposited rapidly. Nannofossils Dominated by cosmopolitan and Tethyan floras, while Boreal were common only in less than one-half the samples examined. floras are absent. However, the assemblages of both sites compare well. In both sites, the Valanginian and Hauterivian are charac- Data available from Neocomian macrofaunas of the Indo- terized by the same index fossils, including C. cuvillieri. In Pacific area suggest that this area is a province of the Tethyan addition to these species, C. salebrosum is common throughout Realm (Mutterlose, this volume). The calcareous nannofossils the Valanginian and lower Hauterivian sequences of both sites. A show a similar pattern. The overall composition of the floras of correlation of the presumably lower Hauterivian and Barremian Sites 765 and 766 suggests moderate-to-weak Tethyan influences sections of both sites is not possible because these strata are barren for the Neocomian. Thus, the Tethyan Realm can be subdivided of nannofossils. In addition, the lower Aptian of Site 765 is also into a Mediterranean Province and an Indo-Pacific Province. barren. In Site 765, the nannofossil-bearing interval starts in the Until Leg 123, little was known about the Neocomian nanno- R. angustus Zone of the late Aptian. However, late Aptian and fossils of the Indo-Pacific area. The nannofloras of the two holes Albian index fossils (R. angustus and P. columnata) are common drilled during Leg 123 are discussed in relation to their paleo- to both sites, allowing for a correlation of these beds. In addition biogeography. to these species, both sites share the same flora. These consist, apart from those species common throughout the Lower Creta- Berriasian-Barremian (Neocomian) ceous, of B. irregularis, F. biforminis, E. varolii, E. apertior, C. Most samples from Holes 765C and 766A yield assemblages litterarius, and C. achylosus. dominated by W. barnesae. Other species found consistently A fairly complete sequence was observed at DSDP Site 261 throughout most of the samples include E.fossacincta, W. britan- about 320 km north of Site 765. The nannofossils at this site, nica, B. constans, B. rotatorius, Zeugrhabdotus embergeri, T. which were described by Proto Decima (1974) and more recently striatum, and L. carniolensis. These species also have consistent by Dumoulin and Bown (this volume), indicate an upper Oxfor- occurrences in both the Tethyan and Boreal realms throughout the dian to Albian age. The Oxfordian age of the oldest sediments in Lower Cretaceous. Thus, these species represent a eurytropic Site 261 is based on Stephanolithion bigotii, a species that is flora. A second group of nannofossils present in the two holes may restricted to the Callovian-Kimmeridgian (Bown et al., 1988). be labeled as Tethyan; however, these only occasionally made The overlying strata yield low-diversity assemblages that consist their way into the Indo-Pacific area. These species include C. of W. manivitae (= C. deflandrei), R. asper, Z. embergeri, W. cuvillieri, S. colligata, C. rothii, Nannoconus spp., and Micran- barnesae, W. communis, C. margerelii, and W. britannica. These tholithus. All these species are rare at both sites, but indicate the have been assigned a Tithonian age (Proto Decima, 1974, p. 595; same distribution pattern for northwestern Europe, where they Dumoulin and Bown, this volume; Table 3). These assemblages also are rare (Mutterlose, 1991). are characterized by the absence of typical Lower Cretaceous A third possible group of species developed independently in species, including L. carniolensis and Cretarhabdus spp. It seems the Indo-Pacific area. Apart from V. matalosa, no other Neo- likely that these Tithonian assemblages, which lack any definite comian species could be referred to in this way. In the Boreal marker species of either a Late Jurassic or age, Realm, V. matalosa has been observed from the latest Barremi- correspond to the low-diversity and low-abundance assemblages an-early Aptian onward; records for the Tethys are not clear. of Site 765 Cores 123-765C-62R-4, -61R-4, and -61R-3 (compare However, this species may be identical to V. compacta, which has

351 Table 2. Distribution of Lower Cretaceous calcareous nannofossils, Hole 766A.

m

Nanno- Vekshinella cf . stradneri Vekshinella stradneri ssp . 1 Vekshinella stradneri ssp . 2 Watznaueria barnesae Watznaueria ovata IRucinolithus sp . Vekshinella quadriarculla Tegumentum octiformis Tubodiscus sp . Vagalapilla matalosa Cretarhabdus crenulatus Tegumentum sp . Watznaueria britannica Grantarhabdus meddii Glaukolithus diplogrammus Chiastozygus tenuis Cretarhabdus conicus Cretarhabdus angustiforatus Cretarhabdus sp . Cretarhabdus surirellus Crucibiscutum salebrosum Cruciellipsis cuvillieri Cyclagelosphaera deflandrei Cyclagelosphaera margerelii Tegumentum striatum Bidiscus rotatorius Micrantholithus (frags. ) Zeugrhabdotus erectus ssp . 1 Zeugrhabdotus sp . Manivitella pemmatoidea Rotelapillus laffittei Assipetra infracretacea Ethmorhabdus hauterivianus Zeugrhabdotus erectus ssp . 2 Biscutum constans Ellipsagelosphaera fossacincta Zeugrhabdotus embergeri Reinhardtites fenestratus Rhagodiscus asper Axoporhabdus dietzmannii plankton servatio n cs r<-> i/-> t- oo O\ o rN m r~~ ON o rN f- oo ON o (N •rt u-> oo ON o 2 rN rN rN rN rN CM m m r<-> r<-> Age zone £ Sample no. •― rN r<-> El 20R-1, 67-68 C R C c R R C R VA F R F F F F R F R R El 20R-2, 103-104 c R F R R F C R R VA F F R R R R F F R R El 20R-3, 95-96 r F F R VA F P r R R R R r p El 20R-4, 35-36 F P F F F F VA R r F R F F F early Albian columnata El 20R, CC F F R F R F VA F F R F R F R R F F El 21R-1, 88-89 F F F R F C F R R VA F R C F R R F F R El 21R-2, 32-34 F C R R R C F F F F F VA F A R R R F C El 21R, CC A R F C F F VA F C C F R R R c A R El 22R, CC r r F R R A F R F F p r p p El 23R-1, 30 r p p r r F F F VA R F R F R . R c R R El 24R-1, 140 F R F R F c F R R VA F R R F R R . R F R R R 1. Aptian- R. angustus El 24R, CC C R R c C R c R F R R VA F F R C R R c R F e. Albian El 25R-1, 55 F F F F r F VA r R p F p El 25R CC F p r r r F p VA p p F El 26R-1, 10 R A R c R R VA C R c R R R R 26R-2, 141-142 early Aptian El 27R-1, 108-109 F R R F R R R R VA R R R R R F E2 27R-3, 18-19 p p p r P p VA R R p R P El 28R-2, 80-81 F R F R R R R F R A F R R R R R R El 28R-4, 62-63 R F R R R VA R R R R F R E2 28R-5, 51-52 F R R F C F R R F C R R A R R F R R F R R R El 28R-6, 69-70 p F R R R R R A R R R R r R Hauterivian- El 29R-1, 119-120 R R F R F R R R A R R R R R R R Barremian E2 29R-3, 4(M1 R F R R R R R R R F R R R R R R R El 29R-4, 17-18 F R R R F R R R R R R R R C R R F R C R E2 30R-1, 14-17 R R R R R R F R R C R R R R R R F R E2 30R-4, 89-92 R R R R F R R R R R R R c R R R R F R R R E2 31R-1, 39-41 R R R F R R R R R c R R R R R R R R R E2 31R-3, 29-31 R R R R R C R F R R c R R R R R R R E2 32R-1,59-60 R R R R R R F R R c R R R R R R F R R E2 32R-2 59-60 r p p p r p p R R p El 32R-4, 48-50 R R R C R R F R c R R R R R R R R E2 3 3R-1,95-96 R R R c R R R R c R R R R R R R R R E2 33R-3, 33-34 R R R R F R R R R R R c R R R R R R R R R R E2 34R-1, 36-38 R R R F R R R R R R R R R R R R R Hauterivian C cuvillieri E2 34R-2, 22-23 R R p p p p R p p F p P R R El 35R-1, 22-24 R R p p F R p p R p R R E2 36R-1, 55-56 p p F R r R F p P p R R E2 36R-2, 17-18 R R R R R R R R R R c R R R R R R R R R R R E2 37R-1, 101-102 R R R R F R R R R R R R R F R R R R R R R R R R R El 37R-3, 28-29 R R R R R R R R F R R R R R R R R R R E2 38R, CC F P R F R R P A R R P F R p R P E2 39R-1, 124-125 R R R R R R R R R F R R R R R Table 2 (continued).

E2 39R-3, 84-85 R R R R R F R F R C R . R R R . R R R E2 40R-1 120 R R F R R R Oi r E2 40R-2, 106 R c R R . R J O J O El 40R-3, 13 R F R R R c R J O R R J O

E2 40R-4 50 R p R X J O X J O J O 5 0 J O F J O R R . R . 5 0 J O 5 0 J O E2 40R-5 76-77 R R R R R p R R R R

O S O S r E2 40R-7, 16-17 R R R c O i S R R R J O J O 5 0 J O E2 41R-2, 29 R R J O J O R c R R R . R J O J O J O R E3 41R-4, 40-41 R F . R R R R Hauterivian C. cuvillieri E3 41R-6, 21-22 F R F R E2 42R-2, 57-59 R R R R R R R R R r R . R R . R R R E2 42R-5, 34-36 R R R R F R R c . R R R . . R R R R E2 43R-2, 82-84 R R R R R R R R r R R R E2 43R-5, 78-80 R R R R R R R R F F R R R . R R E2 44R-1, 129-130 R R F R R R R R R R R R r R R R R R R R E2 44R-3, 118-119 R R R R R R R R F R R R E2 44R-5, 96-97 F R R F R R F R R c R R R R . R . R R R E2 45R-1, 106-108 R R R F R F R R R R r R R R . R R E2 45R-3, 148-149 R R F R F R F R R R c R R R R . R R . R R R ft. L, ft. O S

E2 45R-5, 97-99 R R • O S R c . R R E2 46R-1, 141-142 R F R R R c R R R R J O E2 46R-3, 145-146 R R R R R R F A • J O R J O R R

F.2 46R-5, 138-140 R 5 0 J O R R 5 0 J O R 5 0 J O F A R R X J O R 5 0 J O 5 0 J O • R R O S ft. Pi ft. O S ft, < J O S 47R-1, 80-81 R F R R c R . R R R

47R-3, 51-52 J O F J O c R J O

T i m t n 1 0 t o 47R-5, 50-51 R X 5 0 J O » J O r 5 0 J O 5 0 J O J O R ft. O S U ft, O S ft. ft, ft. O S late E2 47R-7, 29-30 R R R c R R R R R R p P Valanginian- E2 48R-2, 110 R R R C R A R T. striatum early El 48R-3, 10 R R R C C R R R J O Hauterivian E2 48R-4, 140 F R C R J O • A J O R R J O J O J O R R R E2 48R-5, 65-66 F R R r • n j o J O 7 3 J O 5 0 A R p R R R R El 48R-6, 10 R R R R R r R R R R R R r R R R R R R E2 49R-1, 140 R R R r R R R r R R R R . E2 49R-2 120 R R p R R r F R p r R F? 49R-3, 102-108 F R R F E2 49R-4, 58 R p R

c ft. ft. r E2 50R-1, 5 R R R F R R R c R R R R R R R R R A R R

Abundances are characterized by VA (very abundant) = 11-100 specimens/field, A (abundant) = 2-10 specimens/field, C (common) = 1 specimen/1-10 fields, F (few) = 1 specimen/11-100 fields, R (rare) = 1 specimen/more than 100 fields. Preservation: El = slightly etched, E2 = moderately etched, E3 = heavily etched. Table 2 (continued).

Nanno- plankton 1 Prediscosphaera columnata 3 Seribiscutum gaultensis > 8 Flabellites biforaminis > 9 Rotelapillus crenulatus ? 0 Watznaueria biporta '1 Chiastozygus litterarius ' 2 Eprolithus floralis ' 4 Rhagodiscus angustus ' 5 Braarudosphaera hockwoldensis ' 6 Tegumentum stradneri '1 Seribiscutum primitivum ' 8 Crucicribrum striatum constansii ' 9 Parabdolithus infinitus i O Broinsonia signata 5 2 CorolUthion sp . 8 3 Glaukolithus compactus 8 4 Scampanell a cf . cornuta 5 0 Nannoconus sp . 5 7 Rhagodiscus swinnertonnii 5 8 Crucibiscutum cf . salebrosum 5 9 Microstaurius chiastius 6 0 Zeugrhabdotus cf . elegans 6 4 Eprolithus sp . 6 5 Rucinolithus irregularis 6 Grantarhabdus coronadtuentis 6 7 CorolUthion achylosus 8 5 Eprolithus bettenstaedtii Preservatio n 4 Crucibiscutum sp . 4 5 Lithraphidites carniolensis 4 6 Conusphaera rothii 4 7 Diazomatolithus lehmannii 4 9 Chiastozygus sp . 5 1 Haquis circumradiatus 5 2 Tranolithus gabalus 5 4 Sollasites horticus 5 Cretarhabdus schizobrachiatus 5 6 Vekshinella cf . quadriarculla 6 1 Speetonia colligata 6 2 Watznaueria supracretacea 6 3 Cretarhabdus element s Age zone Sample no. 4 2 Hemipodorabdus gorkae 4 3 CorolUthion geometricum 4 8 ISollasites sp . El 20R-1, 67-68 R . . C . . . . . R R . . R . . F R . F C C C R . F F R R R R . R . . R F . F . El 20R-2, 103-104 R C F R R R F C F R . C R . R R R R . R R R R R R R . . R El 20R-3, 95-96 R C R R F R F R C R . C R . F . F C R . F R . R C . . . R P. El 20R-4, 35-36 F R C R F F F F C R C F . . R . R F ...... C . early Albian columnata El 20R, CC R C R F F R C C F C C . . R . R F . R . . R . F . El 21R-1, 88-89 R C R R . R . . F F F F R R F F . R R F C . . . R . R F . El 21R-2, 32-34 C . R . R . F F F F F F F R . F R F R R . . F . F . . . R . El 21R, CC C C R R R F F F C F F . . F R F . . . R R F R F R F . . El 22R, CC F C F F R F F R F R . . R R R R . . . R F R . . El 23R-1.30 c R R . C R . . R . . R R C R . . F R R . . . El 24R-1, 140 R F R F F F . . . F R C R R . R 1. Aptian- R. angustus El 24R, CC C F c R F . R . R R C F e. Albian El 25R-1,55 F F R .FRF F . . C . F F . R . El 25R, CC C F R F R R F C . F . . . F R R R . . El 26R-1, 10 R F R R R R . R R R R 26R-2, 141-142 early Aptian El 27R-1, 108-109 R R R R R R R E2 27R-3, 18-19 R R El 28R-2, 80-81 . . . R R . R . . . El 28R-4, 62-63 . R . E2 28R-5, 51-52 R R R R R late El 28R-6, 69-70 R . R Hauterivian- El 29R-1, 119-120 . . . R . . R R . . R . Barremian E2 29R-3, 40-41 . F . . R . . . . R . . . El 29R-4, 17-18 R R R E2 30R-1, 14-17 R . . R . . . . R . . F . . R R . . . . E2 30R-4, 89-92 . R . R R R R R E2 31R-1, 39-41 R R R R E2 31R-3, 29-31 R R R E2 32R-1,59-60 R R E2 32R-2, 59-60 R R R El 32R-4, 48-50 R R R E2 33R-1, 95-96 R E2 33R-3, 33-34 . R R . R . E2 34R-1, 36-38 R R . R Hauterivian C. cuvillieri E2 34R-2, 22-23 . R . R . . El 35R-1, 22-24 R R E2 36R-1, 55-56 R R R E2 36R-2, 17-18 R R R R E2 37R-1, 101-102 . . . R R . . R . . R F El 37R-3, 28-29 F E2 38R, CC R . . R . E2 39R-1, 124-125 . . . R . Table 2 (continued).

E2 39R-3, 84-85 E2 40R-1, 120 . R . E2 40R-2, 106 El 40R-3, 13 . . . R R E2 40R-4, 50 E2 40R-5, 76-77 R E2 40R-7, 16-17 R E2 41R-2, 29 R R . E3 41R-4, 40^1 . R . Haute rivian C. cuvillieri E3 41R-6, 21-22 . R . E2 42R-2, 57-59 . R . E2 42R-5, 34-36 . R . . R . . . E2 43R-2, 82-84 . . . R . E2 43R-5, 78-80 E2 44R-1, 129-130 . . . R . . . R . . E2 44R-3, 118-119 . R . E2 44R-5, 96-97 R . E2 45R-1, 106-108 . . . R R R . E2 45R-3, 148-149 R R . R . R R . E2 45R-5, 97-99 R R . E2 46R-1, 141-142 . . R R . R . E2 46R-3, 145-146 . . R R R R . E2 46R-5, 138-140 R . R . E2 47R-1, 80-81 R . R R E2 47R-3, 51-52 R R . . . E2 47R-5, 50-51 R . E2 late 47R-7, 29-30 E2 Valanginian- 48R-2, 110 T. striatum El early 48R-3, 10 E2 Hauterivian 48R-4, 140 E2 48R-5, 65-66 El 48R-6, 10 E2 49R-1, 140 E2 49R—2, 120 E2 49R-3, 102-108 E2 49R-4, 58 E2 50R-1, 5 J. MUTTERLOSE

Table 3. Localities where C. salebrosum has been reported (see Fig. 4).

Locality Abundance Time interval Reference

1. Sklinnabanken, Norway Very Ryazanian Crux (1989) abundant 2. Moray Firth, Scotland Abundant Ryazanian- Jakubowski (1987) Barremian Ryazanian- 3. Speeton, England Common Barremian Crux (1989) 4. Northwestern Germany Rare Valanginian— Mutterlose(1991) Aptian 5. DSDP Site 511 Abundant (?)Barremian Wise (1983) 6. Site 692 0.6%-28% Valanginian Mutterlose and Wise (1990) 7. Site 766 Present Valangianian This volume Albian 8. Site 763 Common Valanginian Haq, von Rad, O'Connell, etal. (1990) 9. Site 762 Common Valanginian Haq, von Rad, CTConnell, etal. (1990) 10. Site 761 Common Valanginian Haq, von Rad, O'Connell, et al. (1990) 11. Site 765 Common Valanginian This volume Berriasian- ProtoDecima(1974) 12. DSDP Site 261 Present Albian been mentioned from the Berriasian in Mediterranean sections England). The Speeton Section is located about 1400 km south- (Bralower et al., 1989). So far, V. matalosa is the only species that west of the Norwegian Section. evolved in the Indo-Pacific area. This observation is of impor- This bipolar distribution pattern suggests that C. salebrosum tance because it suggests the origin of southern ocean marine is a stenotropic species characterizing latitudes between 40° and floras. Clarke and Crame (1989) recently discussed the origin of 45°N and 35° to 40°S. Its absence in low latitudes might be southern ocean marine faunas.. explained by temperature control; thus, C. salebrosum may be a Boreal floras (T. septentrionalis, Kokia spp.) and endemic species of moderate-to-cooler waters. The absence of C. salebro- species {Micrantolithus speetonensis) were not observed. The sum in low latitudes may be explained by the following possibili- presence of common C. salebrosum throughout the Valanginian ties: and early Hauterivian at both sites is striking (compare Tables 1 and 2). This species is not present in the late Hauterivian and 1. The species is actually a cool-water morphotype of a tropical Barremian, but reappears in the Aptian. So far, this species has species. not been described from nannofossil-rich Tethyan sections (e.g., 2. The species is present, although extremely rare. Thierstein, 1971, 1973; Applegate et al., 1989; Bralower et al., 3. The species was able to migrate between its two major areas 1989), but is common in the late Berriasian, Valanginian, and late of distribution, the high latitudes, during short periods of cooling. Hauterivian of the Boreal Realm (Mutterlose, 1988, 1991; Crux, 4. The low-latitude forms are represented by an uncalcified, 1989). Figure 4 shows the bipolar distribution of C. salebrosum thus unpreservable, generation. for the Valanginian and early Hauterivian. Sites that contain these species are numbered on the map and listed in Table 3, along with Even if C. salebrosum is described from low-latitudes in the abundances, stratigraphic interval, and references. Based on the future, it is still more common in high latitudes than in low paleogeographic reconstructions by Barron et al. (1981), the lo- latitudes. This distribution is remarkable because no high-latitude calities mentioned here (Sites 261, 511, 765, and 766) had a bipolar distributions have been described so far from the Neo- paleolatitude of 50° to 55°S. However, more recent data by Ogg comian. The absence of Corollithion silvaradion in Holes 765C et al. (this volume) suggest a paleolatitude of 35° to 37°S for Sites and 766A, a species that has been observed both on the Falkland 261,765, and 766 through the Early Cretaceous. Other sites in the Plateau in DSDP Site 511 (Wise, 1983) and in the Boreal Realm southern oceans (DSDP Site 258, ODP Site 693) had a position (Crux, 1989), may be explained by the overall poor preservation of at least 60°S. The northwestern European sites that yield C. of the material studied. salebrosum had a paleolatitude of 40° to 45°N (Barron et al., 1981). Aptian-Albian (Gault) In Holes 765C and 766A, C. salebrosum is common through- The pattern described above changes in the Aptian. A floral out the Valanginian and early Hauterivian interval and declines turnover, linked to a change to open-oceanic conditions, charac- in the middle Hauterivian. Its absence in the early Hauterivian and terizes the early Aptian. This change from a juvenile to a mature Barremian of Hole 765C probably was caused by dissolution. It ocean stage is reflected by the sediments as well (von Rad et al., appears again in the Aptian. In the Boreal Realm, C. salebrosum 1989). A similar turnover was observed in the Boreal Realm is most common in the early Berriasian and Valanginian and (Mutterlose, 1991). This turnover also is characterizing the declines during the early Hauterivian. The species has rarely been macrofaunas of both realms. observed in late Hauterivian and Barremian sequences. Cosmopolitan Neocomian floral components become more Interestingly, in northwestern Europe, this species is most common in the Gault. In addition, several species have their first common in the late Ryazanian, but declines in the Valanginian. occurrence in the early Aptian. These include Rucinolithus irreg- In addition (based on two sites only), an apparent increase in ularis, Chiastozygus litterarius, Flabellites biforaminis, Eprolithus abundance occurs toward higher-latitude sites. Abundances in the floralis, Eprolithus sp., and Seribiscutum gaultensis. Apart from Sklinnabanken Section, which is located off northern Norway, are S. gaultensis, all of these species are known from both the Tethys much higher than those in the Speeton Section (northeastern and Boreal realms. Thus, this Aptian turnover marks the break-

356 LOWER CRETACEOUS NANNOFOSSIL BIOSTRATIGRAPHY

without C. salebrosum with C. salebrosum

SGI-

TETHYAN REALM _

Mediterranean Province

Figure 4. Bipolar distribution of C. salebrosum during the Valanginian and Hauterivian; areas where this taxon is most likely to be found indicated by a stippled pattern. Sites that contain C. salebrosum are numbered on the map and are listed in Table 3, along with abundances (where known), stratigraphic interval, and references. Paleogeographic reconstruction after Barron et al. (1981). < down of Tethyan and Boreal provincialism. The homogenization overlooked in recent studies. However, a bipolar pattern similar of floras may be related to a paleogeographical change, caused by to the one observed for S. primitivum seems another possibility. the breakup of Gondwana and a rise in sea level during the Aptian. However, this idea of a more or less homogeneous flora for the LOWER CRETACEOUS NANNOFOSSIL ZONATION Aptian-Albian interval is modified by two facts: As has already been pointed out, none of the existing nanno- fossil zonations is directly applicable to the Neocomian of the 1. Typical Tethyan genera, such as Nannoconus, Micran- Indo-Pacific area, because all these are based on material from the tholithus, Conusphaera, which are common in the Aptian-Albian Mediterranean area (Thierstein, 1971, 1973; Bralower et al., of the Mediterranean area, are extremely rare in Sites 765 and 766. 1989; Applegate et al., 1989) or on material from the Boreal 2. Both ODP sites yield S. primitivum, which occurs through- Realm (Crux, 1989; Mutterlose, 1988, 1991). Based on the data out the Albian and the Cenomanian. The distribution of this latter gained from Sites 765 and 766, a possible correlation for the species (Fig. 5) has been recently discussed (Roth and Bowdler, Indo-Pacific area, the Mediterranean area, and the Boreal Realm 1981; Roth and Krumbach, 1986; Wise, 1988; Mutterlose and by using calcareous nannofossil events is discussed. Wise, 1990). The species is restricted to high-latitude sites in both the Northern and Southern hemispheres and occurs in two belts Indo-Pacific Nannofloral Events between 35° to 50°N and 35° to 50°S. So far this species has not The nannofossil markers used here to date the Early Creta- been described from low latitudes (compare Fig. 5). This bipolar ceous are based on observations from Sites 765 and 766 (Fig. 6). distribution pattern, comparable to the one shown by C. salebro- The only other data available for the Neocomian of the Indo- sum in the Valanginian and lower Hauterivian intervals, also may Pacific are those from Site 261, which are not detailed enough to be explained by temperature control. Thus, S. primitivum would be integrated. However, additional observations are available for be indicative of a cool surface-water regime, characterizing high- the Aptian-Albian interval (e.g., from the Falkland Plateau [Wise, latitude assemblages. 1983]). Figure 3 shows the locations where Lower Cretaceous rocks were penetrated in the southern ocean. The resolution of the E. ßoralis (= E. apertior) is another species that has been subdivision suggested is limited by the poor preservation within described as a species more common in high-latitudes (Roth and parts of the siliciclastic sequence, in particular, those of late Krumbach, 1986). Not enough data are available for 5. gaultensis; Hauterivian and Barremian age. The following data for the Indo- this fairly small and inconspicuous species may well have been Pacific area have been compiled from Dumoulin and Bown (this

357 J. MUTTERLOSE

o Sites without S. primitivum • Sites with S.primitivum

APTIAN/ALBIAN

Figure 5. Bipolar distribution of S. primitivum during the Aptian to Cenomanian; areas where this taxon is most likely to be found indicated by stippled pattern (from Mutterlose and Wise, 1990). Sites that contain S. primitivum are numbered on the map and are listed in Table 4, along with abundances (where known), stratigraphic interval, and references. Paleogeographic reconstruction after Barron et al. (1981).

Table 4. Localities where S. primitivum has been reported (see Fig. 5).

Locality Abundance Time interval Reference

1. Utah, Wyoming, U.S.A Present Cenomanian Roth and Krumbach (1986) 2. Kansas, U.S.A. Rare Covington(1985) 3. Moray Firth, Scotland Abundant middle Albian Jakubowski(1987) 4. Southern England 0.7%-0.3% early Albian- Roth and Krumbach (1986) late Albian 5. Normandy Coast Present Cenomanian Amedroetal. (1978) Paris Basin Present Albian Manivit(1979) 6. DSDP Site 369 0.2% early Albian Roth and Krumbach (1986) 7. DSDP Site 327 3.8% early Albian Roth and Krumbach (1986) 24.5% late Albian DSDP Site 330 1.6% early Albian Roth and Krumbach (1986) DSDP Site 511 Common early Albian- Wise (1983) 8. Site 693 9.9% late Aptian Mutterlose and Wind (1990) 9. Cauverey Basin Common middle Albian Kale and Phansalkar (1989) 10. Kerguelen Plateau Present Cenomanian- Thierstein (1977) 11. DSDP Site 258 30.0% early Albian Roth and Krumbach (1986) 15.0% late Albian 12. DSDP Site 259 2.6% early Albian Roth and Krumbach (1986) 13. Site 766 Rare late Albian This volume 14. Site 765 Rare late Albian This volume 15. Queensland, Australia Abundant early Albian Shafik (1985)

358 LOWER CRETACEOUS NANNOFOSSIL BIOSTRATIGRAPHY

3. Crucibiscutum salebrosum BOREAL TETHYS INDIAN OCEAN FAD: Berriasian (Site 261) AC: late Valanginian to early Hauterivian (Sites 765, 766) . angustus < 5. Flabellites biforaminis i- < ('FAD F. biforaminis < FAD: early Aptian (Site 766)

IA N IA N FAD: early Aptian (Site 766) Distribution: A cosmopolitan species common in all three LU sLAD C. oblongata LU CC OC areas. OC CC < suggested recently. Based on material from Sites 603 and 638, \NG Applegate and Bergen (1988) and Applegate et al. (1989) devised i < < 12 zones/subzones for the Berriasian-Albian interval (Fig. 6), > / FAD defined by 11 nannofossil events. This zonation is based on the \/W. speetonensis (TAD C. oblongata FADs and LADs of 11 index species. The following datums are used in Fig. 6: Cretarhabdus angustiforatus (FAD late early SIA N / FAD SIA N Berriasian), Calcicalathina oblongata (FAD Berriasian/Valan- \C. angustiforatus ginian), Eiffellithus windii (FAD late early Valanginian), T. stria-

ERRI A ERRI A tum (FAD early Hauterivian), Lithraphidites bollii (FAD early- CO ('FAD C. cuvillieri CO late Hauterivian), S. colligata (FAD early late Hauterivian), C. oblongata(LAD middle Barremian), R. irregularis (FAD early Figure 6. Comparison of different nannofossil zonations for the Boreal Aptian), E. floralis (FAD early late Aptian), and E. turriseiffelii Realm, the Tethys, and the Indo-Pacific area. For the Boreal Realm, the (FAD early Albian). Most of these markers are cosmopolitan zonation is based on Mutterlose (1991), for the Tethys, on Applegate and species that have been observed both in the Boreal Realm and in Bergen (1988), and for the Indian Ocean on this study. FAD = first the Indian Ocean; only L. bollii and C. oblongata are Tethyan appearance datum; LAD = last appearance datum; AC = acme. species. A zonation based on nannofossils has been proposed for the volume) and from this study. The abbreviations used in Figure 6 Boreal Realm recently by Mutterlose (1991). The resolution of are FAD (= first appearance datum), LAD (= last appearance this zonation is much higher than the one for the Tethys (compare datum), and AC (= acme). Fig. 6). However, this zonation is based on Boreal, Tethyan, Boreal endemic, and cosmopolitan species, as well as on mass 1. Cruciellipsis cuvillieri influxes of Tethyan species. In particular, the Valanginian-Bar- FAD: early Berriasian (Site 261) remian interval (Neocomian) is characterized by no cosmopolitan LAD: late Hauterivian (Site 766) markers, while from the Aptian onward, a more homogeneous Distribution: A cosmopolitan species most common in the pattern can be observed. The following datums have been recog- Tethys and also present in both the Boreal Realm and the nized: Micrantholithus speetonensis (LAD late-early Valan- Indo-Pacific area. ginian), T. striatum (FAD early-late Valanginian), C. rothii (FAD 2. Tegumentum striatum early Hauterivian), E. antiquus (FAD early Hauterivian), T. sep- FAD: late Valanginian (Site 765) tentrionalis (FAD late Hauterivian), N. abundans (FAD early LAD: early Hauterivian (Site 766) Barremian), V. matalosa (FAD late Barremian), C. litterarius Distribution: This cosmopolitan species is present in the (FAD early Aptian), E. biforaminis (FAD early Aptian), P. col- Boreal and Tethys realms and the Indo-Pacific area and umnata (FAD early Albian), and E. turriseiffeli (FAD late Al- represents an international marker. However, discrepancies bian). occur in the FAD of this species (early Hauterivian in the In Figure 6,1 try to correlate the zonations of the Boreal Realm, Tethys [Applegate et al., 1989] and early-late Valanginian the Tethys, and the Indo-Pacific area. The FADs and LADs are in the Boreal Realm [Mutterlose, 1991]). similar to the ones described in the text, and the abbreviations

359 J. MUTTERLOSE used correspond to the initials of the species mentioned. From Material. Rare in Site 766, Samples 123-766A-20R-2, 102-104 cm, Figure 6, it becomes obvious that a two-fold zonation exists. and -20R-3, 95-96 cm. While major differences become apparent for the Berriasian-Bar- Diagnosis. A species of Errolithus characterized by 11 regularly remian interval, the Aptian-Albian is characterized by a more or shaped wall elements. Description. Eprolithus bettenstaedtii is round and consists of 11 less homogeneous zonation based on cosmopolitan species. Thus, slightly imbricate vertical wall elements that surround a wide central area. regional zonations have to be used for the Neocomian. However, The width of the wall makes up about one-third of the diameter of the a few cosmopolitan marker species are common in all three areas, whole specimen. The diameter is 5.5 µm. The construction of the central including C. angustiforatus, T. striatum, and C. cuvillieri. S. area is not clear because of poor preservation. colligata is too rare to be considered. On the other hand, a global Differential diagnosis. E. bettenstaedtii differs from contemporane- scheme can be applied for the Aptian-Albian. ous species of the genus Eprolithus (E. apertior, E. varolii) by its 11 wall elements. E. antiquus has about 12 wall elements and is restricted to the SYSTEMATIC PALEONTOLOGY early Hauterivian of the Boreal Realm. Range. F. bettenstaedtii has been observed only from the Albian of Genus SERIBISCUTUM Filewicz et al. in Wise and Wind, 1977 Site 766. Seribiscutum gaultensis n. sp. Geographic distribution. So far, this species has been observed only (PI. l.Figs. 1-6; PI. 6, Fig. 4) from the Indian Ocean. Derivatio nominis. The name is derived from the occurrence of this species in the Aptian-Albian interval (= Gault). Vagalapilla matalosa (Stover, 1966) Thierstein, 1973 Holotype. Plate 1, Figure 5. (PI. 6, Figs. 5-8) Type locality. Site 766: Gascoyne Abyssal Plain. 1966 Coccolithus matalosa Stover, p, 139, pi. 2, figs. 1, 2. Typical stratum. Site 766: Sample 123-766A-21R-2, 32-34 cm: 1973 Vagalapilla matalosa (Stover, 1966) Thierstein, p. 37, pi. 3, figs. 192.82-192.84 mbsf; middle to lower upper Albian, Prediscosphaera 15-18. columnata Zone. Size. Maximum length = 5 µm; maximum width= 3.5 µm, Remarks. V. matalosa differs from other species by a cycle of ele- Material. Consistent occurrence at Site 765, Samples 123-765C-40R- ments inside of the proximal rim. Bars (bridging the central opening in 3, 32-34 cm, to -31R-CC. Consistent occurrence at Site 766, Samples the form of a cross) have typically expanded ends. This species has 123-766A-26R-1, 10 cm, to 20R-1, 67-68 cm. previously been observed only in the upper Barremian-Campanian. At Diagnosis. A small species of Seribiscutum with a central area filled Site 765C, it has its lowest occurrence in the Berriasian-early Valan- by at least four horizontal elements. ginian; at Site 766A, it is consistently present from the early Valanginian Description. Small elliptical placolith constructed of about 20 to 26 to the middle Albian. In the Tethyan and Boreal realms, this species nonimbricate, petaloidal outer elements in the distal shield. The maxi- occurs from the late Barremian/Aptian onward; thus, its presence in the mum length varies between 4 and 5 µm; maximum width is about 3.5 µm. Neocomian of the Indian Ocean may indicate an origin of this species in A second inner distal cycle forms a collar around the central area and is the southern oceans. composed of about 20 vertically, nonimbricate inclined elements. The Known range. Lower Valanginian to Campanian. central area is nearly filled by at least four platelike elements that are Geographic distribution. Worldwide. interrupted in the center by a pore. The central elements are discernible under the light microscope and form a distinctive extinction pattern. ACKNOWLEDGMENTS Differential diagnosis. S. gaultensis differs from B. coronum by its smaller size, its distinctive central elements, and its different stratigraphic I am grateful to J. Bergen (Tulsa, OK), D. K. Watkins (Lincoln, occurrence. 5. gaultensis differs from S. primitivum and S. bijugum, both NE), and an unknown referee for critical reviews and useful of which have plates covering the central area, by its smaller size and the suggestions. Sondra Stewart provided helpful editorial comments. arrangement of its central elements. This species may represent an over- grown Biscutum constans, but it is always discernible from under the light REFERENCES microscope. In addition, the central area of S. gaultensis is wider than Amedro, F., Damotte, R., Manivit, H., Robaszynski, F., and Somay, J., that of B. constans. 1978. Echelles biostratigraphiques dans le Cénomanien du Boulonais Range. S. gaultensis has been observed from the lower Aptian to the (Macro-micro-nannofossiles). Geol. Mediterr., 5:5—18. middle Albian from Sites 765 and 766. Applegate, J. L., and Bergen, J. A., 1988. Cretaceous calcareous nanno- Geographic distribution. So far, this species has been observed only fossil biostratigraphy of sediments recovered from the Galicia Mar- from the Indian and Pacific oceans (Sites 765, 766). gin, ODP Leg 103. In Boillot, G., Winterer, E. L., et al., Proc. ODP, Sci. Results, 103: College Station, TX (Ocean Drilling Program), Crucibiscutum salebrosum (Black, 1971) Jakubowski, 1986 293-348. (PI. 6, Fig.s 1 and 2) Applegate, J. L., Bergen, J. A., Covington, J. M., and Wise, S. S., 1989. 1971 Cruciplacolithus salebrosus Black, p.397, pi.30, fig. 4. Lower Cretaceous calcareous nannofossils from continental margin 1986 Crucibiscutum salebrosum (Black, 1971) Jakubowski, p.38, pi, 1, drill sites off North Carolina (DSDP Leg 93) and Portugal (ODP Leg figs. 16 and 17. 103): a comparison. In Crux, J. A., and Van Heck, S. E. (Eds.), Nannofossils and their Applications: Chichester (Ellis Horwood), Remarks. C. salebrosum has one small central area that is nearly 212-222. completely covered by a massive axial cross constructed of simple solid Barron, E. L, Harrison, C.G.A., Sloan, J. L., and Hay, W. W., 1981. elements. Paleogeography, 180 million ago to the present. Eclogae Geol. Known range. Upper Berriasian/Valanginian to Albian. Helv., 74:443-470. Geographic distribution. Bipolar in high-latitudes. Black, M., 1971. Coccoliths of the Speeton Clay and Sutterby Marl. Proc. Yorks. Geol. Soc, 38:381-424. Genus EPROLITHUS Stover, 1966 , 1973. British Lower Cretaceous coccoliths. I. Gault Clay. 2. Eprolithus bettenstaeditti n. sp. Monogr. Palaeontogr. Soc. London, 127:49-112. (PL 5, Fig. 2; PI. 6, Fig. 19) Black, M., and Barnes, B., 1959. The structure of coccoliths from the Derivatio nominis. Named in honor of the late F. Bettenstaedt for his English Chalk. Geol. Mag., 96:321-328. contributions to studies of Early Cretaceous foraminifers. Bown, P. R., Cooper, M.K.E., and Lord, A. R., 1988. A calcareous Holotype. See Plate 5, Figure 2. nannofossil biozonation scheme for the early to mid Mesozoic. Newsl. Type locality. Site 766, Gascoyne Abyssal Plain. Stratigr., 20:91-114. Typical stratum. Site 766, Sample 123-766A-20R-2, 102-104 cm: Bralower, T. J., 1987. Valanginian to Aptian calcareous nannofossil 183.92-183.94 mbsf. stratigraphy and correlation with the upper M-sequence magnetic Size. Width = 5.5 µm. anomalies. Mar. Micropaleontol., 11:293—310. ,.

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Bralower, T. J, Monechi, S., and Thierstein, H. R., 1989. Calcareous Shafik, S., 1985. Calcareous nannofossils from the , nannofossil zonation of the Jurassic-Cretaceous boundary interval and Eromanga Basin Australia. BMR J. Aust. Geol. Geophys., 9:171-181. correlation with the geomagnetic polarity time scale. Mar. Micro- Sissingh, W., 1977. Biostratigraphy of Cretaceous calcareous nanno- paleontol, 14:153-235. plankton. Geol. Mijnbouw, 56:37-65. Clarke, A., and Crame, J. A., 1989. The origin of the Southern Ocean Stover, L. E., 1966. Cretaceous coccoliths and associated nannofossils marine fauna. In Crame, J. A. (Ed.), Origins and Evolution of the from and the Netherlands. Micropaleontology, 12:133-167. Antarctic Biota. Geol. Soc. Spec. Publ. London, 47:253-268. Thierstein, H. R., 1971. Tentative Lower Cretaceous calcareous nanno- Covington, J. M., 1985. New morphologic information on Cretaceous plankton zonation. Eclogae Geol. Helv., 64:459^88. nannofossils from the Niobrara Formation (Upper Cretaceous) of , 1973. Lower Cretaceous calcareous nannoplankton biostra- Kansas. Geology, 13:683-686. tigraphy. Abh. Geol. Bundesanst. (Austria), 29:1-52. Covington, J. M., and Wise, S. W., Jr., 1987. Calcareous nannofossil , 1977. Mesozoic calcareous nannofossils from the Indian Ocean, biostratigraphy of a Lower Cretaceous deep-sea fan: Deep Sea Drill- DSDP Legs 22 to 27. In Heirtzler, J. R., Bolli, H. M., Davies, T. A., ing Project Leg 93 Site 603, lower continental rise off Cape Hatteras. Saunders, J. B., and Sclater, J. G. (Eds.), Indian Ocean Geology and In van Hinte, J. E., Wise, S. W., Jr., et al , Init. Repts. DSDP, 93: Biostratigraphy. Am. Geophys. Union, 339-351. Washington (U.S. Govt. Printing Office), 617-660. von Rad, U., Thurow, J., Haq, B. U., Gradstein, F., Ludden, J., and ODP Crux, J. A., 1989. Biostratigraphy and palaeogeographical applications Leg 122/123 Shipboard Scientific Parties, 1989. to Cenozoic of Lower Cretaceous nannofossils from north western Europe. In evolution of the NW Australian continental margin and the birth of Crux, J. A., and van Heck, S. E. (Eds.), Nannofossils and Their the Indian Ocean (preliminary results of ODP Legs 122 and 123). Applications: Chichester (Ellis Horwood), 143-211. Geol. Rundsch., 78:1189-1210. Haq, B. U., von Rad, U., O'Connell, S., et al., 1990. Proc. ODP, Init. Wise, S. W., Jr., 1983. Mesozoic and Cenozoic calcareous nannofossils Repts., 122: College Station, TX (Ocean Drilling Program). recovered by Deep Sea Drilling Project Leg 71 in the Falkland Plateau Jakubowski, M., 1986. New calcareous nannofossil taxa from the Lower region, Southwest Atlantic Ocean. In Ludwig, W. J., Krasheninnikov, Cretaceous of the North Sea. INA Newsl., 8:38^12. V. A., et al., Init. Repts. DSDP, 71 (Pt. 2): Washington (U.S. Govt. , 1987. A proposed Lower Cretaceous calcareous nannofossil Printing Office), 481-550. zonation scheme for the Moray Firth Area of the North Sea. Abh. Geol. , 1988. Mesozoic-Cenozoic history of calcareous nannofossils in Bund. (Austria), 39:99-119. the region of the southern ocean. Palaeogeogr., Palaeoclimatol., Kale, A. S., and Phansalkar, V. G., 1989. Black Shale facies from Albian Palaeoecol, 67:157-179. of Cretaceous of South India. Abstr., 28th Int. Geol. Congr., 2:149-150. Wise, S. W., Jr., and Kelts, K. R., 1972. Inferred diagenetic history of a Ludden, J. N., Gradstein, F. M., et al., 1990. Proc. ODP, Init. Repts, 123: weakly silicified deep sea chalk. Trans. Gulf Coast Assoc. Geol. Soc, College Station, TX (Ocean Drilling Program). 22:177-203 Manivit, H., 1979. Le nannoplancton du stratotype de 1'Albien: biozona- Wise, S. W., Jr., and Wind, F. H., 1977. Mesozoic and Cenozoic calcar- tion, systèmatique, nannofaciès, palèoecologie. In Rat, P. (Ed.), L'Al- eous nannofossils recovered by DSDP Leg 36 drilling on the Falkland bian de 1'Aube, Les Stratotypes Francais. C.N.R.S., 307-340. Plateau, Southwest Atlantic sector of the Southern Ocean. In Barker, Manivit, H., Perch-Nielsen, K., Prins, B., and Verbeek, J. W., 1977. Mid P., Dalziel, I.W.D., et al., Init. Repts. DSDP, 36: Washington (U.S. Cretaceous calcareous nannofossil biostratigraphy. Proc. K. Ned. Govt. Printing Office), 269-492. Akad. Wet., Ser. B: Palaeontol., Geol, Phys., Chem., B80:169-181. Zeiss, A., 1983. Zur Frage der Åquivalenz der Stufen Tithon/Berrias/ Mutterlose, J., 1988. Das kalkige Nannoplankton des nw-deutschen Val- Wolga/Portland in Eurasien und Amerika. Ein Beitrag zur Klarung angin und seine palaozeanographische Deutung. Berl. Geowiss. Abh. der weltweiten Korrelation der Jura-/ Kreide-Grenzschichten im Reihe A, 94:175-209. marinen Bereich. Zitteliana, 10:427-438. , 1991. Das Verteilungs- und Migrationsmuster des kalkigen Nannoplanktons in der borealen Unterkreide (Valangin.-Apt.) NW- Date of initial receipt: 24 May 1990 Deutschlands. Palaeontographica B, 221:27-152. Date of acceptance: 16 May 1991 Mutterlose, J., and Wise, S. W., Jr., 1990. Lower Cretaceous nannofossil Ms 123B-124 biostratigraphy of ODP Leg 113 Holes 692B and 693A, continental slope off East Antarctica, Weddell Sea. In Barker, P. F., Kennett, J. P., APPENDIX et al., Proc. ODP, Sci. Results, 113: College Station, TX (Ocean Drilling Program), 325-351. List of species mentioned in the text, arranged in alphabetical Perch-Nielsen, K., 1979. Calcareous nannofossils from the Cretaceous order by generic epithets. between the North Sea and the Mediterranean. In Wiedmann, J. (Ed.), Aspekte der Kreide Europas. IUGS Ser. A, 6:223-272. Assipetra infracretacea (Thierstein, 1973) Roth, 1973 , 1985. Mesozoic calcareous nannofossils. In Bolli, H. M., Saun- Axopodorhabdux dietzmannii (Reinhardt, 1965) Wind and Wise, 1977 ders, J. B., and Perch-Nielsen, K. (Eds.), Plankton Stratigraphy: Bidiscus rotatorius Bukry, 1969 Cambridge (Cambridge Univ. Press), 329-426. Biscutum coronum Wind and Wise in Wise and Wind, 1977 Proto Decima, F., 1974. Leg 27 calcareous nannoplankton. In Veevers, Broinsonia signata Noel, 1969 J. J., Heirtzler, J. R., et al., Init. Repts. DSDP, 27: Washington (U.S. Braarudosphaera hockwoldensis Black, 1973 Govt. Printing Office), 589-621. Calcicalathina oblongata (Worley, 1971) Thierstein, 1971 Roth, P. H., 1978. Jurassic and Lower Cretaceous nannoplankton biostra- Chiastozygus litterarius (Gorka, 1957) Manivit, 1971 tigraphy and oceanography of the northwestern Atlantic Ocean. In Chiastozygus striatum Black, 1971 Benson, W. E., Sheridan, R. E., et al., Init. Repts. DSDP, 44: Wash- Chiastozygus tenuis Black, 1971 ington (U.S. Govt. Printing Office), 731-759. Conusphaera rothii (Thierstein, 1971) Jakubowski, 1986 , 1983. Jurassic and Lower Cretaceous calcareous nannofossils Corollithion achylosus (Stover, 1966) Thierstein, 1971 in the western North Atlantic (Site 534): biostratigraphy, preservation Corollithion geometricum (Gorka, 1957) Manivit, 1971 and some observations on biogeography and paleoceanography. In Corollithion silvaradion Filewicz et al. in Wise and Wind, 1977 Sheridan, R. E., Gradstein, F. M., et al., Init. Repts. DSDP, 76: Cretarhabdus angustiforatus (Black, 1971) Bukry, 1973 Washington (U.S. Govt. Printing Office), 587-621. Cretarhabdus crenulatus Bramlette and Martini, 1964 Roth, P. H., and Bowdler, J., 1981. Middle Cretaceous calcareous nanno- Cretarhabdus schizobrachiatus (Gartner, 1963) Mutterlose, 1991 plankton biogeography and oceanography of the Atlantic Ocean. In Cretarhabdus surirellus (Deflandre in Deflandre and Fert, 1954) Warme, J. E., Douglas, R. G., and Winterer, E. L. (Eds.), The Deep Thierstein, 1971 Sea Drilling Project: a Decade of Progress. Spec. Publ.—Soc. Econ. Cretarhabdus striatus (Stradner, 1963) Black, 1973 Paleontol. Mineral., 32:517-546. Crucibiscutum hayi (Black, 1973) Jakubowski, 1986 Roth, P. H., and Krumbach, K. R., 1986. Middle Cretaceous calcareous Crucibiscutum salebrosum (Black, 1971) Jakubowski, 1986 nannofossil biogeography and preservation in the Atlantic and Indian Crucicribrum striatum constansii Wise and Parker in Wise, 1983 oceans: implications for paleoceanography. Mar. Micropaleontol., Cruciellipsis civillieri (Manivit, 1966) Thierstein, 1971 10:235-266. Cyclagelosphaera deflandrei (Manivit, 1966) Perch-Nielsen, 1985

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Cyclagelosphaera margerelii Noel, 1965 Reinhardtites fenestratus (Worsley, 1971) Thierstein, 1972 Diadorhombus rectus Worsley, 1971 Rhagodiscus angustus (Stradner, 1963) Reinhardt, 1971 Eiffellithus turriseiffelii (Deflandre, 1954) Reinhardt, 1965 Rhagodiscus asper (Stradner, 1963) Reinhardt, 1971 Eiffellithus windii Applegate and Bergen, 1988 Rhagodiscus swinnertonii (Black, 1971) Applegate et al. in Covington Ellipsagelosphaera britannica (Stradner, 1963) Perch-Nielsen, 1968 and Wise, 1987 Ellipsagelosphaera fossacincta Black, 1971 Rotelapillus laffittei (Noel, 1957) Noel, 1973 Ellipsagelosphaera keftalrempti Grün in Grün and Allemann, 1975 Rotelapillus crenulatus (Stover, 1966) Perch-Nielsen, 1984 Eprolithus antiquus Perch-Nielsen, 1979 Rucinolithus irregularis Thierstein in Roth and Thierstein, 1972 Eprolithus bettenstaedtii Mutterlose n. sp. Rucinolithius wisei Thierstein, 1971 Eprolithus floralis (Stradner, 1962) Stover, 1966 Scampanella cornuta Forchheimer and Stradner, 1973 Eprolithus varolii Jakubowski, 1986 Seribiscutum bijugm Filewicz et al. in Wise and Wind, 1977 Etmorhabdus hauterivianus (Black, 1971) Applegate et al. in Coving- Seribiscutum gaultensis Mutterlose n. sp. ton and Wise, 1987 Seribiscutum primitivum (Thierstein, 1974) Filewicz et al. in Wise Flabellites biforaminis Thierstein, 1973 and Wind, 1977 Flabellites oblonga (Bukry, 1969) Crux, 1982 Sollasites horticus (Stradner et al. in Stradner and Adamiker, 1966) Glaukolithus compactus (Bukry, 1969) Perch-Nielsen, 1984 Cepek and Hay, 1969 Glaukolithus diplogrammus Deflandre in Deflandre and Fert, 1954 Speetonia colligata Black, 1971 Grantarhabdus coronadtventis (Reinhardt, 1966) Grün in Grün and Stephanolithion bigotii Deflandre, 1939 Allemann, 1975 Tegumentum octiformis (Köthe, 1981) Crux, 1989 Grantarhabdus meddii Black, 1971 Tegumentum stradneri Thierstein in Roth and Thierstein, 1972 Haqius circumradiatus (Stover, 1966) Roth, 1978 Tegumentum striatum (Black, 1971) Taylor, 1978 Hemipodorhabdus gorkae (Reinhardt, 1969) Grün in Grün and Alle- Tegumentum tripes (Köthe, 1981) Mutterlose, 1988 mann, 1975 Tranolithus gabalus Stover, 1966 Kokia Perch-Nielsen, 1988 Tubodiscus jurapelagicus (Worsley, 1971) Roth, 1973 Lithraphidites bollii (Thierstein, 1971) Theirstein, 1973 Tubodiscus verenae Thierstein, 1973 Lithraphidites carniolensis Deflandre, 1963 Vagalapilla compacta Bukry, 1969 Lithraphidites moray-firthensis Jakubowski, 1986 Vagalapilla matalosa (Stover, 1966) Thierstein, 1973 Manivitella pemmatoidea (Deflandre in Manivit, 1965) Thierstein, Vekshinella quadriarculla (Noel, 1965) Rood et al., 1971 1971 Vekshinella stradneri ssp. 1 Rood et al., 1971 Micrantholithus Deflandre in Deflandre and Fert, 1954 Vekshinella stradneri ssp. 2 Micrantholithus hoschulzii (Reinhardt, 1966) Thierstein, 1971 Watznaueria barnesae (Black in Black and Barnes, 1959) Perch- Micrantholithus obtusus Stradner, 1963 Nielsen, 1968 Micrantholithus speetonensis Perch-Nielsen, 1979 Watznaueria biporta Bukry, 1969 Microstaurus chiastius (Worsley, 1971) Grün in Grün and Allemann, Watznaueria britannica (Stradner, 1963) Reinhardt, 1964 1975 Watznaueria communis Reinhardt, 1964 Microstaurus quadratus Black, 1971 Watznaueria manivitae (Forchheimer, 1972) Proto Decima, 1974 Nannoconus Kamptner, 1931 Watznaueria ovata Bukry, 1969 Nannoconus abundans Stradner and Grün, 1973 Watznaueria supracretacea (Reinhardt, 1965) Wind and Wise, 1976 Nannoconus globulus Brönnimann, 1955 Zeugrhabdotus embergeri (Noel, 1959) Perch-Nielsen, 1984 Nannoconus steinmannii Kamptner, 1931 Zeugrhabdotus elegans Gartner, 1968 Parabdolithus infinitus (Worsley, 1971) Thierstein in Roth and Thier- Zeugrhabdotus erectus ssp. 2 stein, 1972 Zeugrhabdotus erectus ssp. 1 Prediscosphaera columnata (Stover, 1966) Perch-Nielsen, 1984 Zeugrhabdotus sisyphus (Gartner, 1968) Crux, 1989

362 LOWER CRETACEOUS NANNOFOSSIL BIOSTRATIGRAPHY

Plate 1. Biscutaceae and Ahmuellerellaceae from Hole 766A. All specimens from Sample 123-766A-21R-2, 32-34 cm. Scale of the bars of the photomicrographs = lµm. 1-6. Seribiscutum gaultensis n. sp. 7-9. Vekshinella stradneri Rood et al., D.

363 J. MUTTERLOSE

Plate 2. Ahmuellerellaceae, Chiastozygaceae, Ellipsagelosphaeraceae, and Podorhabdaceae from Hole 766A. Figure 6 is from Sample 123-766A-20R-2, 103-104 cm; all other figures from Sample 123-766A-21R-2, 32-34 cm. Scale of bars = 1 µm. 1, 2. Broinsonia signata (Noel); (1) P, (2) D. 3, 4. Vekshinella stradneri Rood et al.; P. 5, 7. Chiastozygus cf. litterarius (Gorka); (5) P, (7) D. 6. Flabellites biforaminis Thierstein; P. 8, 9. Manivitella pemmatoidea (Deflandre in Manivit); P.

364 LOWER CRETACEOUS NANNOFOSSIL BIOSTRATIGRAPHY

Plate 3. Ellipsagelosphaeraceae, Podorhabdaceae, and Polycyclolithaceae from Hole 766A. Figures 1 through 7 from Sample 123-766A-21R-2, 32-43 cm; Figures 8 and 9 from Sample 123-766A-20R-2, 102-104 cm. Scale bar = 1 µm. 1. Watznaueria britannica (Stradner); D. 2. Cretarhabdus conicus Bramlette and Martini; D. 3, 6, 7. Rucinolithus irregularis (Thierstein in Roth and Thierstein). 4, 5. Grantarhabdus coronoadtventis (Reinhardt); (4) P, (5) D. 8. Eprolithus sp. 9. Eprolithus floralis (Stradner).

365 J. MUTTERLOSE

Plate 4. Polycyclolithaceae, Rhagodiscaceae, and Stephanolithiaceae from Hole 766A. Figure 1 from Sample 123-766A-20R-2, 102-104 cm; all other figures from Sample 123-766A-21R-2, 32-34 cm. Scale bar = 1 µm. 1. Eprolithus bettenstaedtii n. sp. 2, 3. Eprolithus floralis (Stradner). 4, 5. Rhagodiscus angustus (Stradner); (4) P; (5) D. 6-8. Rotelapillus crenulatus (Stover); (6, 7) side views; (8) P. 9. Rotelapillus laffittei (Noel); P.

366 LOWER CRETACEOUS NANNOFOSSIL BIOSTRATIGRAPHY

Plate 5. Stephanolithiaceae, Zygodiscaceae, and inc. sedis from Hole 766A and light photomicrographs. Figures 1 through 4, and 6 from Sample 123-766A-21R-1, 32-34 cm; Figure 5 from Sample 123-766A-20-2, 102-104 cm. Scale bar = 1 µm. 1-3. Corollithion achylosus Stover; (1, 3) D: (2) P. 4, 5. Glaukolithus diplogrammus (Deflandre in Deflandre and Fert); (4) D; (5) P. 6. Scampanella cf. cornuta Forchheimer and Stradner (Figs. 7-11 2700×. 7,8. Cyclagelosphaera deflandrei (Manivit); oval form; Sample 123-765C-62R-1, 120 cm; (7) TL; (8) POL. 9. Cruciellipsis cuvilliere (Manivit); Sample 123-765C-56R-2, 60-61 cm; POL. 10. Speetonia colligata Black; Sample 123-765C-56R-2, 60-61; POL. 11. Corollithion achylosus (Stover); Sample 123-766A-20R-4, 35-36 cm; POL.

367 J. MUTTERLOSE

17

2 3

Plate 6. Photomicrographs from Holes 765C and 766A. Figure 24, 1800×, all other figures, 2700×. 1, 2. Crucibiscutum salebrosum (Black); POL; (1 Sample 123-765C-56R-2, 102-104 cm; (2) Sample 123-766A-48R-4, 140 cm. 3. Seribiscutum primitivum (Thierstein); POL; Sample 123-766A-20R-2, 103-104 cm. 4. Seribiscutum gaultensis n. sp.; POL; Sample 123-766A-20R-1, 67-68 cm. 5-8. Vagalapilla matalosa (Stover); (5) Sample 123-766A- 48R-4, 140 cm; (6-8) Sample 123-766A-50R-1, 5 cm; (5, 6, 8) POL; (7) TL. 9-11. Tegumentum striatum (Black); POL; (9) Sample 123-766A-50R-1, 5 cm; (10, 11) Sample 123-766A-48R-4, 140 cm. 12-15. Flabellites biforaminis Thierstein; POL; (12, 13) Sample 123-766A-20R-4, 35-36 cm; (14, 15) Sample 123-766A-20R-1, 67-68 cm. 16. Chiastozygus litterarius (Gorka); POL; Sample 123-766A-20R-1, 67-68 cm. 17. Eprolithus floralis (Stradner); POL; Sample 123-766A-20R-1, 67-68 cm; 18. Eprolithus sp.; POL; Sample 123-766A-20R-1, 67-68 cm. 19. Eprolithus bettenstaedtii n. sp.; POL; Sample 123-766A-20R-3, 95-96 cm. 20-23. Rucinolithus irregularis Thierstein; (20, 22) POL; (21, 23) TL; (20, 21) Sample 123-766A- 21R-CC. 24. Rhagodiscus angustus (Stradner); POL; Sample 123-765C-39R-4, 30-31 cm. 25. Prediscosphaera spinosa (Bramlette and Martini); POL; Sample 123-766A-20R-3, 95-96 cm.

368