UPPER ALBIAN AMMONITES FROM ODP LEG 171B OFF NORTHERN FLORIDA

by JENS LEHMANN

ABSTRACT. ODP Leg 171B investigated the sediments of the Blake Plateau off northern Florida and recovered 36 Upper Albian ammonites – one from Site 1050C, the others from Site 1052E. This unusually large number of specimens from an ODP site permits the dating of the interval between 668 to 621 m below sea-floor at Site 1052E as late Late Albian, Stoliczkaia (S.) dispar ammonite zone. This zone is indicated by the genera Mortoniceras and Stoliczkaia (S.). Site 1050C (Interval 171B-1050C-31R-3, 0·80–0·86 m) cannot be dated more precisely than Late Aptian to Mid Cenomanian by ammonites. The fauna is cosmopolitan. Tetragonites jurinianus and Puzosia mayoriana are widely distributed forms. Kossmatella muhlenbecki was thought to be restricted to a fairly small area around the Mediterranean, but the record off northern Florida presented here, indicates that it is not an endemic species; this is also true for Hemiptychoceras subgaultinum in the Albian. The event-like character of the ammonite-bearing interval at Site 1052E is unique. It is overlain by a laminated claystone succession; the top of this sequence is considered to represent maximum flooding (Oceanic Anoxic Event, OAE 1d). Ammonites perhaps profited from an increased nutrient supply derived from flooded coastal plains during a continuous transgression.

C RETACEOUS ammonites from DSDP and ODP material have rarely been described (e.g. Wiedmann and Neugebauer 1978; Renz 1983; Young 1984) because they are generally too widely scattered to be recovered in borehole (drill) cores. In February 1997, several ammonites were recovered during ODP Leg 171B at the edge of the Blake Plateau off northern Florida, in the western Atlantic (Shipboard Scientific Party 1998a, p. 1052, fig. 14; Text-fig. 1). I resampled the ammonite-bearing interval at the Bremen Core Repository (BCR), Germany in December 1997. Thirty-five specimens were recovered from Cores 1052E- 57R–51R [668–621 m below sea floor (mbsf)]. The assemblage indicates the Upper Albian, upper Stoliczkaia (S.) dispar Zone and provides important new stratigraphical data, as well as data on ammonite palaeobiogeography and palaeoenvironmental conditions.

PRESERVATION In ODP material, often only the aptychi of ammonites are preserved (e.g. Renz 1979) because the Mesozoic rocks at many sites were deposited below the aragonite compensation depth. Consequently, the common occurrence of ammonites in the Upper Albian at Site 1052E (Text-fig. 1) is striking and the event- like character of their occurrence has to be explained. All of the ammonites from Site 1052E are crushed and commonly flattened, and their original nacreous shell is preserved. This indicates deposition above the aragonite compensation depth. In some cases, internal moulds are coated by pyrite. The single fragment (Sample 171B-1050C-31R-3W, 0·80–0·86 m) from Site 1050C is a three- dimensional, pyritic steinkern with remains of the shell attached. This preservation also indicates deposition above the aragonite compensation depth. All specimens are preserved in the collections of the Bremen Core Repository, Cores 1052E-57R–51R (668–621 mbsf) and 1050C-31R-3W, 0·80–0·86 m. Only material that is of palaeogeographical and/or stratigraphical importance is included in the Systematic Palaeontology section, but other less significant taxa are shown in Table 1 and Text-figure 2.

[Palaeontology, Vol. 43, Part 1, 2000, pp. 41–61, 1 pl.] ᭧ The Palaeontological Association 42 PALAEONTOLOGY, VOLUME 43

TEXT-FIG. 1. Sketch map showing the ODP Leg 171B Blake Nose drilling area in the western North Atlantic with position of transect drill sites and the multichannel seismic line (MCS Line) TD 5. Ammonites have been sampled from sites 1050 and 1052; spots NE of Site 1052 are adjacent Sites 1053 and 1051, spot NE of Site 1050 is Site 1049 (DSDP 390). Modified after Shipboard Scientific Party (1998b, figs 1, 20).

STRATIGRAPHICAL IMPLICATIONS The present material represents a typical late Late Albian ammonite fauna and undoubtedly can be referred to the Stoliczkaia (S.) dispar Zone of the European ammonite zonal scheme (e.g. Owen 1984; Hancock 1991). The ammonite assemblage shows similarities to middle and upper S.(S.) dispar Zone (respectively the Mortoniceras (Durnovarites) perinflatum Subzone and Arrhaphoceras (Praeschloenbachia) briacen- sis Subzone) faunas of the proposed stratotype for the Albian–Cenomanian boundary at Mont Risou, south-east France (Gale et al. 1996). In fact, with the exception of Tetragonites jurinianus, all ammonites from Site 1052E that have been determined at species level (Kossmatella muhlenbecki, Puzosia mayoriana and Hemiptychoceras subgaultinum also occur at Mont Risou). There is no evidence of the older subzone of M.(M.) rostratum. A correlation with previously described ammonite faunas from DSDP Leg 40 off western Africa (Wiedmann and Neugebauer 1978) is not possible. These included the Upper Albian ammonites Puzosia quenstedti (Parona and Bonarelli, 1897), Cainoceras sp. nov. ex aff. liberum van Hoepen, 1942 and Puzosia mayoriana (d’Orbigny, 1841). Only the last-named is reported herein from Leg 171B. Young (1984) assigned faunas from Sites 535 and 540 of DSDP Leg 77 in the Gulf of Mexico to the LEHMANN: UPPER ALBIAN AMMONITES 43

TEXT-FIG. 2. Ammonite occurrences at ODP Leg 171B, Site 1052E. C, Cenomanian; Ss, Substage; N, nannofossil zone; F, foraminifer zone; AZ, ammonite zone; AS, ammonite subzone. Lithological section modified after Shipboard Scientific Party (1998a, fig. 1).

Upper Albian, ‘Stoliczkaia dispar and/or Arrhaphoceras substuderi zones’. These faunas, including Hypophylloceras cf. guillantoni (Collignon, 1932), an indeterminate lytoceratid, Stomohamites cf. virgulatus (Brongniart, 1822), Turrilitoides sp. juv. cf. T. toucasi (He´bert and Munier-Chalmas, 1875), and Scaphites sp. nov. aff. S. simplex Jukes-Browne, 1875, are dominated by heteromorphs, and have no genera in common with Leg 171B. Integration of ammonite, microfossil and nannofossil stratigraphy in the Upper Albian is unsatisfactory on a global scale. Mont Risou is the best section for comparison with the present data, since macrofossils, microfossils and calcareous nannofossils (Gale et al. 1996) of a large part of the Upper Albian S. (S.) dispar Zone there have been investigated. Correlation of planktonic foraminiferal events at Site 1052E, Leg 171B (data from Shipboard Scientific Party 1998c, Table 7) and the Mont Risou succession (shown in Text-fig. 4) indicates that the present ammonite fauna is from the M.(D.) perinflatum Subzone rather than the A.(P.) briacensis Subzone. 44 PALAEONTOLOGY, VOLUME 43

TABLE 1. Ammonites from ODP Leg 171B, sites 1052E and 1050C.

Material from Leg 171B, site 1052E Core Sc cm no. of specimens

Hamites cf. duplicatus 51 1 50 or 54 1 Anagaudryceras ? 52 2 51 or 53 1 Desmoceras cf. latidorsatum 52 1 114–116 1 Kossmatella sp. 53 5 80 1 Puzosia mayoriana 53 4 94 1 Zelandites cf. odiense 53 4 91 1 Hemiptychoceras subgaultinum 53 3 60 1 Puzosia mayoriana 53 2 48 1 Tetragonites sp. 53 1 126 1 Kossmatella muhlenbecki 53 1 93 1 Hamites ? 53 1 23 1 Hamites ? 54 2 126 or 127 1 Hemiptychoceras subgaultinum 54 CC* 40–41 1 ammonite, indet. 55 6 63 1 Hemiptychoceras subgaultinum 55 6 36 1 Zelandites ? 55 6 25 1 ammonite, indet. 55 5 96 1 Puzosia ? 55 5 95 1 Lechites ? 55 4 104 1 lytoceratid ammonit, indet. 55 4 87·5 1 ammonite, indet. 55 64 85·5 2 Puzosia ? 55 4 85·5 1 Stoliczkaia (Stoliczkaia) sp. 55 4 62 1 Tetragonites ? 55 4 48 1 Puzosia mayoriana 55 4 30 1 Phyllopachyceras sp. 55 4 0–2·5 1 Puzosia sp. juv. 55 3 140 1 Tetragonites jurinianus 55 3 c. 62 1 Mortoniceras sp. 55 3 59·5 2 Puzosia sp. juv. 55 2 10 1 Kossmatella muhlenbecki 55 1 141 1 Lechites ? 55 1 132 1 Desmoceras ? 55 1 117·5 1 Hemiptychoceras subgaultinum 57 6 73 1

Material from Leg 171B, site 1050C

Phylloceras sp. 31 3W 80–86 1 *CC ¼ core catcher

This interpretation, based on correlation of planktonic foraminifera events, can be confirmed by calcareous nannofossils (Text-Fig. 4). In the Mont Risou section, the boundary between nannofossil subzones CC9a and 9b and that between M.(D.) perinflatum and A.(P.) briacensis ammonite subzones are almost coincident (Text-fig. 4; Gale et al. 1996, fig. 2). According to the nannofossil data at Site 1052E (Shipboard Scientific Party 1998a, fig. 20 and Text-fig. 2), this boundary is more than 90 m above the top of the ammonite-bearing interval (Text-fig. 4). Although ranges of planktonic foraminifera and calcareous nannofossils are comparable between the proposed stratotype at Mont Risou and Site 1052E, integration of the planktonic foraminiferal and calcareous nannofossil stratigraphy is still problematic. According to Bralower et al. (1995, fig. 7), the first occurrence of the foraminiferal zonal marker LEHMANN: UPPER ALBIAN AMMONITES 45 Biticinella breggiensis correlates with the boundary between calcareous nannofossil zones CC9a and 9b but this species already occurs at 682·57 m below sea floor at Site 1052E, a level which is believed to represent calcareous nannofossil zone CC8b (Shipboard Scientific Party 1998c, table 7 and fig. 20). The single ammonite recovered from Site 1050C of Leg 171B (Table 1; sample 171B-1050C-31R-3W, 0·80–0·86 m) is identified as Phylloceras sp. of the long-ranging Suborder Phylloceratina. According to planktonic foraminifera, this sample comes from the Upper Albian, Rotalipora ticinensis Zone (Shipboard Scientific Party 1998c, p. 121, fig. 29). If the interval (605·39–577·3 m below sea floor) characterized by common R. ticinensis at Site 1050C (Shipboard Scientific Party 1998c, p. 126) and at Mont Risou correspond (Gale et al. 1996, fig. 7), the horizon of the specimen is older than, or as old as, the lower Arrhaphoceras (Praeschloenbachia) briacensis Subzone (Stoliczkaia (S.) dispar Zone). However, the occurrence of R. ticinensis is stated to be ‘rare’ (lowermost sample) and ‘few’ (all other samples) in table 10 of Shipboard Scientific Party 1998c (p. 127) in contrast to the statement on p. 126.

PALAEOBIOGEOGRAPHY The opening of seaways by sea-floor spreading in the Atlantic led to a major global transgression in the mid Cretaceous. This is supposed to be the main reason for the generally observed, increasingly cosmopolitan character of ammonite faunas at the very end of the Early Cretaceous (Wiedmann 1988). Rising sea level eliminated geographical barriers, although it has been shown that sea-level rise probably also led to provincialism at that time (Marcinowski and Wiedmann 1988). Ammonites, as mainly bottom-related animals, strongly reacted to a reduction of shallow- water conditions and associated climatic changes. However, Klinger and Wiedmann (1983) pointed out that our understanding of the geographical distribution of Cretaceous ammonites merely reflects the state of our present knowledge and that just a few additional records could change the picture entirely. The general composition of the fauna from ODP Leg 171B is Tethyan. This can be seen from the comparatively high proportion of ammonites belonging to the Lytoceratina (30 per cent.) and Phyllocer- atina (3 per cent.; cf. Systematic Palaeontology and Table 1). However, the fauna is dominated by representatives of the Suborder Ammonitina (nearly 40 per cent.) and a high proportion (c. 30 per cent.) of Ancyloceratina. Some of the species recorded from Leg 171B are widely distributed forms (Tetragonites jurinianus, Puzosia mayoriana), although there are very few data on Albian ammonoid occurrences in the North Atlantic (Text-fig. 3). In contrast, Kossmatella muhlenbecki was thought to be restricted to a fairly small area around the Mediterranean but its record from the slope off northern Florida presented here indicates that it is not an endemic species. In the Albian, Hemiptychoceras subgaultinum was known previously from south-west Europe and Hungary only (Text-fig. 3). The present record is again evidence for a distinctly wider distribution. However, this is not surprising because Cenomanian representatives of this species have been recorded in Madagascar, far from its main area of distribution (Text-fig. 3).

PALAEOENVIRONMENT More than 60 per cent. of the Leg 171B material consists of leiostracan (smooth shelled) ammonites, most of which are lytoceratids, phylloceratids and desmoceratids. In general, the proportion of leiostracan ammonites is known to increase with depth (e.g. Westermann 1996) and, according to several authors, an oceanic deep-water character would therefore be indicated for the fauna from Leg 171B (see review by Westermann 1996). Deposition on the middle or lower slope is therefore assumed for the Upper Albian at the site. The acme-occurrence of ammonites is directly overlain by a laminated claystone succession (Text-figs 46 PALAEONTOLOGY, VOLUME 43

TEXT-FIG. 3. Palaeobiogeographical distribution of ammonite species found at ODP Leg 171B using the plate tectonic reconstruction for the latest Albian (100 My) of Hay et al. (in press). Dots indicate occurrence of species; stars indicate Cenomanian records (where distinction is made between Albian and Cenomanian records). LEHMANN: UPPER ALBIAN AMMONITES 47

TEXT-FIG. 3 cont. 48 PALAEONTOLOGY, VOLUME 43

TEXT-FIG. 4. Biostratigraphical markers of the Upper Albian, Stoliczkaia (S.) dispar Zone of Leg 171B, Site 1052E and the proposed stratotype of the Albian-Cenomanian boundary at Mont Risou near Rosans, south-east France. Arrows against the stratigraphical column for Site 1052E indicate position of Albian benthic foraminiferal fauna. Note that the Albian-Cenomanian boundary determined by Shipboard Scientific Party (1998a) appears to be too low at Site 1052E compared with the Risou section. Lithology of Site 1052E modified after Shipboard Scientific Party (1998a, fig. 1); for legend, see Text-fig. 2. Mont Risou section after Gale et al. (1997); note that lowest foraminiferal sample obtained from there is at ¹136 m.

2, 4), the very top part of which has yielded a poor benthic foraminiferal fauna (Shipboard Scientific Party 1998a, p. 267). All other Albian samples are comparatively rich in these forms (Text-fig. 4). The poor foraminiferal fauna is presumed to reflect Oceanic Anoxic Event 1d (OAE 1d; Erbacher and Thurow 1997). This agrees with the correlation of biostratigraphical events between Leg 171B, Site 1052E and the well-controlled section at Mont Risou in south-east France (Text-fig. 4). OAE 1d, called the Breitroffer level in the Vocontian Basin, comprises an interval c. 5 m above and below the boundary between calcareous nannofossil subzones CC9a and 9b, but although this boundary has not been determined with certainty at Site 1052E, it fits perfectly with the position of the poor benthic foraminiferal fauna. Deposition of the laminated claystone sequence preceded the productivity event of OAE 1d; the latter is related to maximum sea level, leading to a higher nutrient supply from flooded coastal plains (P-OAE; Erbacher et al., 1996). Ammonites probably took advantage of the rising sea level and the associated increased primary production of zoo- and phytoplankton that started before the maximum flooding event, but unfortunately, nothing is known about the ammonite fauna of the laminated claystone interval, so it is only possible to speculate. LEHMANN: UPPER ALBIAN AMMONITES 49

SYSTEMATIC PALAEONTOLOGY

Order AMMONOIDEA von Zittel, 1884 Suborder PHYLLOCERATINA Arkell, 1950 Superfamily PHYLLOCERATACEAE von Zittel, 1884 Family PHYLLOCERATIDAE von Zittel, 1884 Subfamily PHYLLOCERATINAE von Zittel, 1884 Genus PHYLLOCERAS Suess, 1865

Phylloceras sp. Plate 1, figures 6–7

Material. Sample 171B-1050C-31R-3W, 0·80–0·86 m preserved as a fragment of a steinkern with remains of the shell. This is the only ammonite sample obtained from core 1050C. Remarks. The well-preserved fragmentary steinkern shows part of the upper flank and venter (maximum whorl breadth c. 7·2 mm, whorl height c. 6·75 mm). It has fine, straight, radial ribs on the venter. Its flanks are nearly smooth, flattened, subparallel, as in early whorls of Phylloceras (Hyporbulites) seresitense Wiedmann, 1964, Phylloceras (H.) subseresitense Pervinquie`re, 1907, Phylloceras (H.) aptiense applanatum Wiedmann, 1964 and Phylloceras (Phylloceras) serum Oppel, 1865 (see Wiedmann 1964, and Kennedy and Klinger 1977 for descriptions). A specimen determined as ‘Hypophylloceras sp. cf. H. tanit’ by Young (1979, fig. 9h) also has subparallel flanks, but not before a whorl height of c. 40 mm. Among phylloceratids with very flattened subparallel flanks, Phylloceras (Hyporbulites) seresitense shows straight or slightly adapical bent ribs across the venter at an early growth stage like the present specimen. However, the latter cannot be determined at species level because of its fragmentary preservation and the inadequate description in the literature of the earliest whorls of Phylloceras species. The ribbing on the venter is similar to that of several species of Gaudryceras de Grossouvre, 1894 and Anagaudryceras Shimizu, 1934, but at the early growth stage represented by sample 171B-1050C-31R- 3W, 0·80–0·86 m, these genera are characteristically smooth, with the exception of constrictions, or have proportionally much finer ventral ribs that are usually clearly projected (Kennedy and Klinger 1979, Matsumoto 1995). Furthermore, early whorls of these two genera are inflated to rather compressed and do not show clearly flattened, almost parallel, flanks like the present specimen. Some early whorls of the Albian genus Zuluscaphites Van Hoepen, 1955 also have straight and fine ribs across the venter (e.g. Z. orycteropusi Van Hoepen, 1955 figured by Kennedy and Delamette 1994, fig. 13–3); however, the inner flanks of these forms are inflated and the outer flanks are convergent at this stage in contrast to the present specimen. Furthermore, Zuluscaphites is ventrally wedge-shaped and not subrectangular; its flanks are flat only on the body-chamber (see diagnosis of genus by Kennedy and Klinger 1993, p. 64; Kennedy and Delamette 1994, figs 12–13).

Distribution. Sample 171B-1050C-31R-3W, 0·80–0·86 m comes from c. 600 m below the sea floor at Site 1050C and is dated as Late Albian, Rotalipora ticinensis Zone of the planktonic foraminiferal zonation and CC9a Subzone of the calcareous nannoplankton stratigraphy (Shipboard Scientific Party 1998c, p. 121, fig. 29).

Genus PHYLLOPACHYCERAS Spath, 1925a

Phyllopachyceras sp.

Text-figure 5A

Material. Sample 171B-1052E-55R-4, 0·000–0·025 m. 50 PALAEONTOLOGY, VOLUME 43 Remarks. The present specimen, with a maximum diameter of c. 29 mm, is poorly preserved but the few features observed are typical of the genus. The umbilicus is tiny; on the inner flanks, there are prorsiradiate, very fine striae whereas the outer flank is ornamented with radial, strong but blunt, broad ribs, that are as broad as the intercostal spaces. The specimen is crushed and the whorl section appears highly compressed. After photographing the specimen (Text-fig. 5A), a part of the inner flank of the body- chamber was removed to expose the suture lines (features of the removed part of the shell surface are still visible on the counterpart). It shows the typical tetraphyllic endings on the first and second lateral saddle. Ornamentation, supposed whorl section and probable type of umbilicus are very similar to those of an undescribed specimen of about the same size from California, USA. The latter is a well-preserved phragmocone with a crushed body-chamber, showing remains of the shell (University of California, Riverside, coll. Murphy; UCR no. 379/2, ex coll. UC Los Angeles 3015 from the Upper Aptian, Gabbioceras wintunium Zone, WSW of Ono, Shasta County, California, locality 3015 of Murphy 1956, fig. 4). The ribbing of the present specimen is similar to that of Phylloceras shastalense Anderson, 1902, a further form from California, mature shells of which are of comparable size but show an inflated whorl section with depressed whorls (Anderson 1902, pl. 4, fig. 114; 1938, pl. 12, fig. 6). The type and all additional material of P. shastalense Anderson was collected at Horsetown, Shasta County, California and is Early Albian in age (M. A. Murphy, pers. comm. 1998).

Distribution. The present specimen comes from the upper part of the Upper Albian (Text-fig. 2).

Suborder LYTOCERATINA Hyatt, 1889 Superfamily TETRAGONITACEAE Hyatt, 1900 Family TETRAGONITIDAE Hyatt, 1900 Genus TETRAGONITES Kossmat, 1895

Tetragonites jurinianus (Pictet, 1847)

Plate 1, figure 5

*1847 Ammonites Jurinianus Pictet, p. 297, pl. 3, fig. 3. 1988 Tetragonites jurinianus (Pictet, 1847); Delanoy and Latil, pl. 1, fig. 2a–c. 1990 Tetragonites jurinianus (Pictet); Marcinowski and Wiedmann, p. 32, pl. 2, fig. 1; text-fig. 18 [see for further synonymy]. 1997 Tetragonites jurinianus (Pictet); Delamette et al., pl. 40, fig. 7.

Material. Sample 171B-1052E-53R-3, 0·62 m.

Remarks. The present specimen is well preserved with a maximum diameter of c. 12 mm. It has a well- rounded whorl section and no ornament. Specimens with a rounded whorl section and a lack of constrictions throughout their ontogeny have been conventionally assigned to T. jurinianus. Scholz (1979) included the latter in the highly variable species T. timotheanus (Pictet, 1847) but later authors (e.g. Wright and Kennedy 1984, 1996; Marcinowski and Wiedmann 1990; Gale et al. 1996) have been able to work only on small collections of Tetragonites, and a discussion of Scholz’s (1979) ideas is still needed.

Distribution. Representatives of Tetragonites occur from the uppermost Aptian to the Maastrichtian. T. jurinianus occurs in the Upper Albian of south-west Europe, Africa and Central America (Text-fig. 3). LEHMANN: UPPER ALBIAN AMMONITES 51

TEXT-FIG.5.A, Phyllopachyceras sp., sample 171B-1052E-55R-4, 0·000–0·025 m. B, Stoliczkaia (Stoliczkaia) sp., sample 171B-1052E-55R-4, 0·62 m. Both × 2.

Family GAUDRYCERATIDAE Spath, 1927 Genus ANAGAUDRYCERAS Shimizu, 1934

Anagaudryceras? sp. Plate 1, figure 11

Material. Sample 171B-1052E-52R-2, 0·51 or 0·53 m.

Remarks. Although the fragment (maximum length c. 36 mm) is diagenetically distorted, the coiling is serpenticone. The specimen appears entirely smooth, but the shell surface is badly damaged. Since there is no sign of ornament, the material is better placed in Anagaudryceras Shimizu, 1934 than in Gaudryceras de Grossouvre, 1894. Gaudryceras is similar to Anagaudryceras but has coarser ribs and lirae (Wright 1996, p. 3). Its serpenticone whorls distinguish it from the moderately evolute coiling of Tetragonites. The coiling is similar to that of the tetragonitid Saghalinites Wright and Matsumoto, 1954, which is serpenticone but with slightly depressed whorls, and which is known from the Santonian to Maastrichtian only (Wright 1996, p. 7).

Distribution. The genus Anagaudryceras ranges from the Middle Albian to the Maastrichtian and occurs in England, France, Germany, Austria, Romania, Algeria, Angola, South Africa, Madagascar, India, Japan, Sakhalin, Kamchatka, Alaska, British Columbia, California, New Zealand and Antarctica. 52 PALAEONTOLOGY, VOLUME 43

Genus ZELANDITES Marshall, 1926

Zelandites cf. odiensis (Kossmat, 1895) Plate 1, figure 8 cf. *1895 Lytoceras (Gaudryceras) Odiense sp. nov. Kossmat, p. 129 (33), pl. 58 (4), fig. 1a–c. cf. 1979 Zelandites odiensis (Kossmat); Kennedy and Klinger, p. 163, pl. 14, fig. 4 [see for further synonymy].

Material. Sample 171B-1052E-53R-4, 0·91 m.

Remarks. The present specimen has a diameter of c. 14·5 mm. It is poorly preserved but the involute coiling, compressed whorl section and lack of any ornament indicate that it probably belongs to Z. odiensis (Kossmat, 1895); see Kennedy and Klinger (1979) for comparison with related taxa.

Distribution. Zelandites odiensis is known from the Albian of Madagascar and the Cenomanian of India and South Africa.

Genus KOSSMATELLA Jacob, 1907

Kossmatella muhlenbecki (Fallot, 1885) Plate 1, figures 2, 17 *1885 Ammonites muhlenbecki Fallot, p. 233, pl. 4, fig. 1. 1996 Kossmatella (Kossmatella) muhlenbecki (Fallot); Gale et al., p. 551, fig. 10a, c–e [see for further synonymy].

Material. Samples 171B-1052E-53R-1, 0·93 m; 171B-1052E-55R-1, 1·41 m.

Remarks. Both specimens are moderately evolute and show a fine dense ribbing throughout ontogeny. The only additional ornament comprises typical radial folds, first developing at a whorl height of c. 5·5 mm in

EXPLANATION OF PLATE 1 Fig. 1. Puzosia mayoriana (d’Orbigny, 1841); sample 171B-1052E-53R-4, 0·94 m; × 2. Fig. 2. Kossmatella muhlenbecki (Fallot, 1885); sample 171B-1052E-53R-1, 0·93 m; × 2. Fig. 3. Lechites?; sample 171B-1052E-55R-4, 1·04 m; × 2. Fig. 4. Hamites cf. duplicatus Pictet and Campiche, 1861; sample 171B-1052E-51R-1, 0·50 or 0·54 m ; × 2. Fig. 5. Tetragonites jurinianus (Pictet, 1847); sample 171B-1052E-55R-3, c. 0·62 m; × 1. Figs 6–7. Phylloceras sp.; sample 171B-1050C-31R-3W, 0·80–0·86 m. 6, ventral view. 7, right lateral side. Both × 2. Fig. 8. Zelandites cf. odiensis (Kossmat, 1895); sample 171B-1052E-53R-4, 0·91 m; × 2. Fig. 9. Puzosia mayoriana (d’Orbigny, 1841); sample 171B-1052E-53R-2, 0·48 m; × 2. Fig. 10. Hemiptychoceras subgaultinum Breistroffer, 1940; sample 171B-1052E-54R-CC, 0·40–0·41 m; × 2. Fig. 11. Anagaudryceras?; sample 171B-1052E-52R-2, 0·51 or 0·53 m; × 1. Fig. 12. Hemiptychoceras subgaultinum Breistroffer, 1940; sample 171B-1052E-55R-6, 0·36 m; × 2. Figs 13–14. Desmoceras cf. latidorsatum (Michelin, 1838); sample 171B-1052E-52R-1; 1·14–1·16 m. 13, right lateral side; note oblique distortion. 14, left lateral side. Both × 1. Figs 15–16. Hemiptychoceras subgaultinum Breistroffer, 1940. 15, sample 171B-1052E-53R-3, 0·6 m. 16, sample 171B-1052E-57R-6, 0·73 m. Both × 2. Fig. 17. Kossmatella muhlenbecki (Fallot, 1885); sample 171B-1052E-55R-1, 1·41 m; × 2. Fig. 18. Mortoniceras sp., two specimens, sample 171B-1052E-55R-3, 0·595 m; × 2. PLATE 1

LEHMANN, ammonites 54 PALAEONTOLOGY, VOLUME 43 sample 171B-1052E-53R-1, 0·93 m which has a maximum diameter of c. 14 mm. The larger specimen (171B-1052E-55R-1, 1·41 m; maximum diameter c. 29 mm) shows 14 folds per whorl (at a diameter of c. 28 mm). The ribs are narrow and distinct on the first three-quarters of the last whorl and broader and less prominent on the last quarter. Sample 171B-1052E-53R-1, 0·93 m shows c. 7 folds on the last whorl; it represents an early growth stage that has not yet developed the folds for one whole whorl. The fine and dense riblets on the shell of the present specimens are comparable to those seen on specimen 1315/20 of the Geologisch-Pala¨ontologisches Institut und Museum der Universita¨t Tu¨bingen (Wiedmann and Dieni 1968) which has recently been re-examined. The latter specimen shows that an early smooth stage is seen only on steinkerns. The 14 folds per whorl of 171B-1052E- 55R-1 correspond well with the number given by Gale et al. (1996). The lack of tubercles on the umbilical edge and lack of a bullatiforme intermediate stage, in addition to the fairly distinct folds and moderately evolute whorls, distinguish this species from other taxa of the genus (cf. Wiedmann and Dieni 1968).

Distribution. K. muhlenbecki is known from the Albian of southern France, Spain, Sardinia, and Hungary (Text-fig. 3).

Suborder AMMONITINA Hyatt, 1889 Superfamily DESMOCERATACEAE von Zittel, 1884 Family DESMOCERATIDAE von Zittel, 1895 Genus PUZOSIA Bayle, 1878

Puzosia mayoriana (d’Orbigny, 1841) Plate 1, figures 1, 9 *1841 Ammonites mayorianus d’Orbigny, p. 267, pl. 79, figs 1–3. 1990 Puzosia mayoriana mayoriana (d’Orbigny); Marcinowski and Wiedmann, p. 55, pl. 5, fig. 5. 1990 Puzosia mayoriana provincialis (Parona and Bonarelli); Marcinowski and Wiedmann, p. 57, pl. 6, fig. 3; fig. 25a–b. 1998b Puzosia mayoriana (d’Orbigny, 1841); Lehmann, p. 407, figs 1a–d, 2a–d [see for further synonymy]. 1998 Puzosia mayoriana (d’Orbigny, 1841); Kaplan et al., p. 71, pl. 1, figs 12–13; pl. 9, figs 1, 11–12; pl. 10, figs 3–5 [see for further synonymy]. 1998 Epipuzosia europaea Lehmann, 1994; Kaplan et al., p. 74, pl. 3, fig. 35. 1998 Parapuzosia (Austiniceras) austeni (Sharpe, 1855); Kaplan et al., p. 78, pl. 1, figs 4, 7, 9; pl. 2; pl. 4.

Material. Samples 171B-1052E-53R-4, 0·94 m; 171B-1052E-53R-2, 0·48 m.

Remarks. P. mayoriana (d’Orbigny, 1841) is a highly variable species. In recent years, it has been discussed extensively (e.g. Cooper and Kennedy 1987; Marcinowski and Wiedmann 1990; Lehmann 1998b) but the major revision called for by Marcinowski and Wiedmann (1990) has not yet been achieved. Both the present specimens show distinct and flexuous constrictions, and therefore can be easily distinguished from representatives of the Puzosia quenstedti group. Sample 171B-1052E-53R-4, 0·94 m has a diameter of c. 11·5 mm and 171B-1052E-53R-2, 0·48 m, a maximum length of c. 29 mm. Without new and extensive collections, further discussion on the relationships of species of this genus is pointless; P. provincialis is therefore included in the synonymy only provisionally.

Distribution. P. mayoriana (in the sense used here) ranges from the Middle Albian to the Upper Cenomanian in Europe, Africa, India, and Japan (Text-fig. 3; compare Wright and Kennedy 1984).

Genus DESMOCERAS von Zittel, 1884 LEHMANN: UPPER ALBIAN AMMONITES 55 Desmoceras cf. latidorsatum (Michelin, 1838) Plate 1, figures 13–14 cf. *1838 Ammonites latidorsatus Nobis Michelin, p. 101, pl. 12, fig. 9. cf. 1994 Desmoceras (Desmoceras) latidorsatum (Michelin, 1838); Kennedy, p. 219, pl. 2, figs 13–14; pl. 4, figs 14–17; pl. 6, figs 7–8. cf. 1996 Desmoceras (Desmoceras) latidorsatum (Michelin, 1838); Gale et al., p. 551, figs 11h–j, 13d, o, 171 (pars). cf. 1997 Desmoceras latidorsatum (Michelin); Delamette et al., pl. 13, fig. 8; pl. 18, fig. 1.

Material. Sample 171B-1052E-52R-1, 1·14–1·16 m. Remarks. D. latidorsatum is a highly variable species (e.g. Marcinowski and Wiedmann 1990). The present specimen shows distinct and strongly curved constrictions, as in the specimens figured by Renz (1982, pl. 6, figs 3a–b). Although the ODP material is strongly and obliquely distorted, the whorl section can be envisaged as inflated but it cannot be determined whether it is as inflated as in the inflatum morphotype (Breistroffer 1933).

Distribution. D. latidorsatum has a world-wide distribution and is recorded from the Lower Albian to Upper Cenomanian.

Superfamily ACANTHOCERATACEAE de Grossouvre, 1894 Family MOJSISOVICSIIDAE Hyatt, 1903 Subfamily MORTONICERATINAE Douville´, 1912 Genus MORTONICERAS Meek, 1876

Mortoniceras sp. Plate 1, figure 18

Material. Sample 171B-1052E-55R-3, 0·595 m (two specimens on one bedding plane). Remarks. The smaller specimen is incompletely preserved, with a maximum diameter of c. 10 mm, but clearly shows a smooth keel, distinct umbilical bullae, and associated or intercalated primary and secondary ribs. Some of the ribs bear bullae somewhat above the mid-flank. These features do not allow the species to be recognized. The larger specimen, with a maximum diameter of c. 20·8 mm, has smooth innermost whorls except for a few umbilical bulges. After this, there are distinct, sharply curved (?flexuous) ribs. The venter of the outer whorl shows a smooth keel. Any further details of the ornament are not clearly recognizable, and it is therefore not possible to assign the specimen to any previously described species.

Distribution. The genus ranges from the upper Middle Albian to the uppermost Upper Albian and is known from several European countries, Africa, India, North and South America, and Madagascar.

Family LYELLICERATIDAE Spath, 1921 Genus STOLICZKAIA Neumayr, 1875

Stoliczkaia (Stoliczkaia) sp.

Text-figure 5B

Material. Sample 171B-1052E-55R-4, 0·62 m is an internal mould (illustrated); a rubber cast of the external mould shows part of the shell preserved. 56 PALAEONTOLOGY, VOLUME 43

Remarks. The internal mould (Text-fig. 5B) is poorly preserved (partly worn and pyritized) with a maximum diameter of c. 39 mm. It shows radial ribs that become prorsiradiate on the outer flank. The ribs are fairly dense (c. 14 per quarter whorl), somewhat narrower than the interspaces, and distinct but not prominent. Primary ribs are present across the whole flank and secondaries intercalated on the outer half of the flank. Parts of the original shell are still visible on the external mould, but most of the shell adhered to the rubber cast. The latter indicates that the mode of ornament on the shell is different from that visible on the steinkern, with both well-rounded ribs on the inner half of the whorl and only narrow intercostal areas. The last part of the outer whorl shows that the sculpture becomes more prominent and distant, with well separated ribs; it then becomes smooth with coarse growth lines on the last part of the shell, indicating maturity at an estimated diameter of 50 mm. It could therefore represent a microconch. I hesitate to assign this limited material to any species, because ontogenetic changes and intraspecific variability have not been investigated in any great detail.

Distribution. The subgenus Stoliczkaia first appears in the lower part of the Stoliczkaia (S.) dispar Zone of the Upper Albian and ranges throughout the zone (Wright and Kennedy 1994, p. 550, fig. 1). It is widely distributed in Europe and also occurs in the Crimea, Ukraine, Turkmenia, Tunisia, Nigeria, Angola, Madagascar, southern India, Texas, Mexico, Brazil and Japan.

Suborder ANCYLOCERATINA Wiedmann, 1966b Superfamily TURRILITACEAE Gill, 1871 Family HAMITIDAE Gill, 1871 Genus HAMITES Parkinson, 1811

Hamites cf. duplicatus Pictet and Campiche, 1861 Plate 1, figure 4 cf.* 1861 Hamites duplicatus Pictet and Campiche, p. 98. cf. 1998a Hamites duplicatus Pictet and Campiche; Lehmann, p. 29, pl. 5, fig. 3. cf. 1998 Hamites duplicatus Pictet and Campiche, 1861; Kaplan et al., p. 174, pl. 58, figs 1–3; pl. 59, fig. 2 [see for further synonymy].

Material. Sample 171B-1052E-51R-1, 0·50 or 0·54 m.

Remarks. The flattened specimen has a length of c. 15 mm. It shows fine, more or less rectiradiate, dense ribbing, c. 11 per whorl height suggesting H. duplicatus Pictet and Campiche, 1861. However, apart from the poor preservation of this specimen, the general problem of separating this taxon from other species is well known, as recently pointed out by Wright and Kennedy (1995).

Distribution. The cosmopolitan genus Hamites ranges from the Lower Albian to the Upper Cenomanian. Albian records of H. duplicatus are known from England, Switzerland and France. Cenomanian representatives occur in England, Poland, Daghestan, Mangyshlak and Kazakhstan.

Genus HEMIPTYCHOCERAS Spath, 1925b

Hemiptychoceras subgaultinum Breistroffer, 1940 Plate 1, figures 10, 12, 15–16 *1940 Hemiptychoceras subgaultinum Breistroffer, p. 159. 1996 Hemiptychoceras subgaultinum Breistroffer; Gale et al., p. 569, figs 21a–e, g–i, k; 22b, e–g; 23b [see for further synonymy]. LEHMANN: UPPER ALBIAN AMMONITES 57

Material. Samples 171B-1052E-54R-CC, 0·40–0·41 m; 171B-1052E-53R-3, 0·6 m; 171B-1052E-55R-6, 0·36 m; 171B-1052E-57R-6, 0·73 m.

Remarks. Samples 171B-1052E-55R-6, 0·36 m (Pl. 1, fig. 12; c. 16 mm long), 171B-1052E-57R-6, 0·73 m (Pl. 1, fig. 16; c. 21 mm long) and 171B-1052E-54R-CC, 0·40–0·41 m (Pl. 1, fig. 10; c. 20 mm long) show 8–10 fine and sharp ribs per whorl height, although the specimens are crushed. This number indicates the high rib density (six or seven ribs per whorl height) typical for this species. Furthermore, the frequency of constrictions in this material and in 171B-1052E-53R-3, 0·6 m (Pl. 1, fig. 15; c. 22·5 mm long) corresponds to the material recently figured by Gale et al. (1996) from south-east France.

Distribution. H. subgaultinum is known from the upper Upper Albian of south-west Europe and Hungary, the Cenomanian of the Crimea and Madagascar, and the lowermost Cenomanian of southern France.

Family BACULITIDAE Gill, 1871 Genus LECHITES Nowak, 1908

Lechites? sp. Plate 1, figure 3

Material. Two fragments: samples 171B-1052E-55R-1m, 1·32 m; 171B-1052E-55R-4, 1·04 m.

Remarks. The specimens are nearly smooth; only the figured fragment 171B-1052E-55R-4, 1·04 m (16 mm long, 4·5 mm wide) shows a single rib at one end. They could be representatives of the near-smooth or very sparsely ornamented end forms of Lechites gaudini Pictet and Campiche, 1861. This variant has been described by Breistroffer (1947, p. 62) as raricostata and was figured by Wiedmann and Dieni (1968, pl. 6, fig. 5a–b) and Gale et al. (1996, fig. 27f). Like the present specimens, French material from the Upper Albian of Mount Risou near Rosans, housed at the Geologisch-Pala¨ontologisches Institut, Tu¨bingen, Germany (no registration numbers), shows that flattened juveniles of this variant are unornamented.

Distribution. The genus Lechites Nowak, 1908 ranges from the Upper Albian to the Lower Cenomanian. Like L. gaudini from the Upper Albian S. (S.) dispar Zone and its equivalents, it has been described from several European countries, North and South Africa, Madagascar, India, Japan and Mexico.

Acknowledgements. I thank J. Erbacher (Hannover) and T. Pletsch (Kiel) for obtaining the original ammonite samples during ODP Leg 171B and for their general comments, help and reviews. W. Gerber (Tu¨bingen) took the photographs and K. Piolka (Twist) made additional prints. I am indebted to M. A. Murphy (University of California, Davis) for fruitful discussions and assistance; M. Kirby (also Davis) kindly corrected the English. I gratefully acknowledge the comments made by two anonymous referees. The samples described herein have been collected for the Sonder- forschungsbereich 275 ‘Klima’ of the DFG (German Research Foundation) at Tu¨bingen. This publication was supported by a grant of the common university special program 3 of the Federal Republic of Germany and the single states of the Federal Republic provided by the DAAD (German Academic Exchange Program).

REFERENCES

ANDERSON, F. M. 1902. Cretaceous deposits of the Pacific Coast. Proceedings of the California Academy of Science, Third Series, Geology, 1, 1–155. —— 1938. Lower Cretaceous deposits in California and Oregon. Memoirs of the Geological Society of America, Special Papers, 16, 1–244. ARKELL, W. J. 1950. A classification of the Jurassic ammonites. Journal of Paleontology, 24, 354–364. BAYLE, E´ . 1878. Fossiles principaux des terrains. Explication de la Carte Ge´ologique de France, Service de la Carte Ge´ologique Detaille´e, Paris, 4, 1–158. 58 PALAEONTOLOGY, VOLUME 43

BRALOWER, T. J., LECKIE, M. R., SLITER, W. V. and THIERSTEIN, H. R. 1995. An integrated Cretaceous microfossil biostratigraphy. Special Publication of the Society for Sedimentary Geology (SEPM), 54, 65–79. BREISTROFFER, M. 1933. Etudes de l’e´tage albien dans le massif de la Chartreiuse (Ise`re et Savoie). Traveaux de Laboratoire Ge´ologique de Faculte´ Scientifique de l’Universite´ Grenoble, 17, 1–50. —— 1940. Re´vision des ammonites du Vraconien de Salazac (Gard) et conside´rations ge´ne´rales sur ce sous-e´tage albien. Traveaux de Laboratoire Ge´ologique de Faculte´ Scientifique de l’Universite´ Grenoble, 22 (for 1938–39), 71–171. —— 1947. Sur les zones d’ammonites dans l’Albien de France et d’Angleterre. Travaux de Laboratoire Ge´ologique de Faculte´ Scientifique de l’Universite´ Grenoble, 26 (1946–47), 17–104. BRONGNIART, A. 1822. Sur quelques terrains de Crai hors du Bassin de Paris. 80–101. In CUVIER, G. and BRONGNIART, A. Description ge´ologiques des environs de Paris. 3rd edition. Dufour et D’cagne, Paris, 428 pp. COLLIGNON, M. 1932. Les ammonites pyriteuses de l’Albien supe´rieur du Mont Raynaud a´ Madagascar. Annales Geologiques du Service des Mines, Madagascar, 2, 2–36. COOPER, M. R. and KENNEDY, W. J. 1979. Uppermost Albian (Stoliczkaia dispar Zone) ammonites from the Angolan littoral. Annals of the South African Museum, 77, 175–308. —— —— 1987. A revision of the Puzosiinae (Cretaceous ammonites) of the Cambridge Greensand. Neues Jahrbuch fu¨r Geologie und Pala¨ontologie, Abhandlungen, 174, 105–121. DELAMETTE, M., CHAROLLAIS, J., DECROUEZ, D. and CARON, M. 1997. Les gre`s verts helve´tiques (Aptien moyen-Albien superieur) de Haute-Savoie, Valais et Vaud (Alpes Occidentales Franco-Suisses). Analyse stratigraphique et inventaire pale´ontologique. Publication du De´partement de Ge´ologie et Pale´ontologie, 23, i–xi þ 1–400. DELANOY, G. and LATIL, J.-L. 1988. De´couverte d’un nouveau gisment Albien dans les environs de Drap (Alpes- Maritimes, France) et description d’une riche ammonitofaune d’age Albien terminal. Ge´obios, 21, 749–771. DOUVILLE´ ,H.1912. E´ volution et classification des Pulchelliide´s. Bulletin de la Societe´ Ge´ologique de France, 11, 285– 320. ERBACHER, J. and THUROW, J. 1997. Influence of oceanic anoxic events on the evolution of mid-Cretaceous radiolaria in the North Atlantic and western Tethys. Marine Micropaleontology, 30, 139–158. —— —— and LITTKE, R. 1996. Evolution patterns of radiolaria and organic matter variations: a new approach to identify sea-level changes in mid-Cretaceous pelagic environments. Geology, 24, 499–502. FALLOT, E. 1885. E´ tudes ge´ologique sur les e´tages moyens et supe´rieurs du terrain Cre´tace´ dans le sud-est de la France. Annales Scientifique de Ge´ologie, Paris, 18, 1–263. GALE, A. S., KENNEDY, W. J., BURNETT, J. A., CARON, M. and KIDD, B. E. 1996. The Late Albian to Early Cenomanian succession at Mont Risou near Rosans (Droˆme, SE France): an integrated study (ammonites, inoceramids, planktonic foraminifera, nannofossils, oxygen and carbon isotopes). Cretaceous Research, 17, 515–606. GILL, T. 1871. Arrangement of the families of mollusks. Smithsonian Miscellaneous Collections, 227, i–xvi þ 1–49. GROSSOUVRE, A. de 1894. Recherches sur la Craie Supe´rieur. 2, Pale´ontologie. Les ammonites de la Craie Supe´rieure. Me´moires de la Service de Carte De´taille Ge´ologique de France, 264 pp. HANCOCK, J. M. 1991. Ammonite scales for the Cretaceous System. Cretaceous Research, 12, 259–291. HAY, W., DECONTO, R., WOLD, C. N., WILSON, K. M., VOIGT, S., SCHULZ, M., WOLD-ROSSBY, A., DULLO, W.-C., RONOV, A. B., BALUKHOVSKY, N. and SOEDING, E. in press. An alternative global Cretaceous paleogeography. In JOHNSON, E. and BARRERA, C. (eds). The evolution of the Cretaceous ocean/climate systems. Special Papers of the Geological Society of America. HE´ BERT, E. and MUNIER-CHALMAS, E. C. P. A. 1875. Appendice pale´ontologique. Description du terrain supe´rieur de Bassin d’Uchaux. Annales des Sciences Geologique, 6, 113–152. HOEPEN, E. C. N. van 1942. Die gekielde Ammoniete van die Suid-Afrikaanse Gault. 2. Drepanoceratidae, Pervinquier- idae, Arestoceratidae, Cainoceratidae. Paleontologiese Navorsing van die Nasionale Museum Bloemfontein, 1,91– 157. —— 1955. New and little-known ammonites from the Albian of Zululand. South African Journal of Science, 51, 355– 361 HYATT, A. 1889. Genesis of the Arietidae. Smithonian Contributions of Knowledge, 673, i–xi þ 1–239. —— 1900. Cephalopoda. 502–604. In ZITTEL, K. A. von (ed.). Textbook of palaeontology, 1896–1900. Macmillan Press, London and New York, 1438 pp. —— 1903. Pseudoceratites of the Cretaceous. Monograph of the United States Geological Survey, 44, 1–250. JACOB, C. 1907. E´ tudes pale´ontologiques et stratigraphiques sur la partie moyenne des terrains cre´tace´s dans les Alpes franc¸aise et les regions voisines. Annales de l’Universite´ Grenoble, 19, 221–534. JUKES-BROWNE, A. J. 1875. On the relations of the Cambridge Gault and Greensand. Quarterly Journal of the Geological Society, London, 31, 256–314. KAPLAN, U., KENNEDY, W. J., LEHMANN, J. and MARCINOWSKI, R. 1998. Stratigraphie und Ammonitenfaunen des westfa¨lischen Cenoman. Geologie und Pala¨ontologie in Westfalen, 51, 1–236. LEHMANN: UPPER ALBIAN AMMONITES 59

KENNEDY, W. J. 1994. Cenomanian ammonites from Cassis, Bouches-du-Rhoˆne, France. 209–254. In Proceedings of the 3rd Pergola International Symposium, Palaeopelagos. Special Publication, 1, Pergola, 420 pp. —— and DELAMETTE, M. 1994. Lyelliceratidae and Flickiidae (Ammonoidea) from the Upper Albian of the Helvetic Shelf (Western Alps, France and Switzerland). Journal of Paleontology, 68, 1263–1284. —— and KLINGER, H. C. 1977. Cretaceous faunas from Zululand and Natal, South Africa. The ammonite family Phylloceratidae. Bulletin of the British Museum (Natural History), Geology Series, 27, 347–380. —— —— 1979. Cretaceous faunas from Zululand and Natal, South Africa. The ammonite family Gaudryceratidae. Bulletin of the British Museum (Natural History), Geology Series, 31, 121–174. —— —— 1993. On the affinities of Zuluscaphites Van Hoepen 1955 (Cretaceous Ammonoidea) from the Albian of Zululand, South Africa. Pala¨ontologische Zeitschrift, 67, 63–67. KLINGER, H. C. and WIEDMANN, J. 1983. Palaeobiogeographic affinities of Upper Cretaceous ammonites of northern Germany. Zitteliana, 10, 413–425. KOSSMAT, F. 1895. Untersuchungen u¨ber die su¨dindische Kreideformation. Beitra¨ge zur Pala¨ontologie und Geologie von O¨ sterreich-Ungarn, 9, 97–203. LEHMANN, J. 1994. The ammonite Pachydesmoceras aff. denisonianum (Stoliczka) described by O. Renz 1976 from the Lower Cenomanian of Cressier, Switzerland is reassigned to Epipuzosia europaea n. sp. Neues Jahrbuch fu¨r Geologie und Pala¨ontologie, Monatshefte, 7, 407–417. —— 1998a. Systematic palaeontology of the ammonites of the Cenomanian-Lower Turonian of northern Westphalia, North Germany. Tu¨binger Geowissenschaftliche Arbeiten, Reihe A, 37, 1–58. —— 1998b. Taxonomy and nomenclature of large desmoceratids (Ammonoidea) from the Cenomanian and Turonian of Western Europe. Neues Jahrbuch fu¨r Geologie und Pala¨ontologie, Abhandlungen, 207, 405–418. MAEDA, H. and SEILACHER, A. 1996. Ammonoid taphonomy. 543–578. In LANDMAN, N., TANABE, K. and DAVIS, R. A. (eds). Ammonoid paleobiology. Topics in Geobiology, 13. Plenum Press, New York, 857 pp. MARCINOWSKI, R. and WIEDMANN, J. 1988. Paleogeographic implications of the Albian ammonite faunas of Poland. 491– 504. In WIEDMANN, J. (ed.). Cephalopods – present and past. Schweizerbart, Stuttgart, 965 pp. —— 1990. The Albian ammonites of Poland. Palaeontologia Polonica, 50, 1–94. MARSHALL, P. 1926. The Upper Cretaceous ammonites of New Zealand. Transactions of the New Zealand Institute, 56, 129–210. MATSUMOTO, T. 1995. Note on gaudryceratid ammonites from Hokkaido and Sakhalin. Special Papers of the Palaeontographical Society of Japan, 35, 1–152. MEEK, F. B. 1876. A report on the invertebrate Cretaceous and Tertiary fossils of the upper Missouri country. In HAYDEN, F. V. (ed.). Reports of the United States Geological Survey of the Territories, 9, i–lxiv þ 1–629. MICHELIN, H. 1838. Note sur une argile de´pendant du Gault. Me´moires de la Socie´te´ Ge´ologique de France, 3, 97–103. MURPHY, M. 1956. Lower Cretaceous stratigraphic units of Northern California. Bulletin of the American Association of Petroleum Geologists, 40, 2098–2119. NEUMAYR, M. 1875. Die Ammoniten der Kreide und die Systematik der Ammonitiden. Zeitschrift der Deutschen Geologischen Gesellschaft, 27, 854–942. NOWAK, J. 1908. Untersuchungen u¨ber Cephalopoden der Oberen Kreide in Polen. I. Teil, Genus Baculites. Bulletin International de l’Academie des Sciences de Cracovie, (Series B), 1908, 326–353. OPPEL, A. 1865. Die tithonische Etage. Zeitschrift der Deutschen Geologischen Gesellschaft, 17, 535–558. ORBIGNY, A. C. V. D. d’ 1841. Pale´ontologie francaise: Terrains cre´tace´s. 1. Cephalopodes. Masson, Paris, 121–430. OWEN, H. G. 1984. The Albian stage: European province chronology and ammonite zonation. Cretaceous Research, 5, 329–344. PARKINSON, J. 1811. Organic remains of a former world. J. Robson, London, 479 pp. PARONA, C. F. and BONARELLI, G. 1897. Fossili albiani d’Escragnolles, del Nizzardo e della Liguria occidentale. Palaeontographia Italica, 2, 53–112. PERVINQUIE` RE, L. 1907. Carte ge´ologique Tunisie. E´ tudes de pale´ontologie tunisienne. 1, Cephalopodes des terrains secondaires. Paris, i–v þ 438 pp. PICTET, F. J. 1847. De´scription des mollusques fossiles qui se trouvent dans les Gre`s Verts des environs de Gene`ve. Me´moires de la Socie´te´ de Physique et d’Histoire Naturelle de Gene`ve, 11, 257–412. —— and CAMPICHE, G. 1861–1864. Mate´riaux pour la Pale´ontologie Suisse, 2, Description des fossiles du terrain cre´tace´ des environs de Sainte-Croix. Gene`ve, 752 pp. RENZ, O. 1979. Aptychi (Ammonoidea) and ammonites from the Lower Cretaceous of the Western Bermuda Rise, Leg 43, Site 387, DSDP. 591–597. In TUCHOLKE, B. E., et al. (eds). Initial Reports of the Deep Sea Drilling Project, 43, 1115 pp. —— 1982. The Cretaceous ammonites of Venezuela. Birkha¨user, Basel, 132 pp. —— 1983. Early Cretaceous cephalopoda from the Blake-Bahama Basin (Deep Sea Drilling Project Leg 76, Hole 60 PALAEONTOLOGY, VOLUME 43

534 A) and their correlation in the Atlantic and southwestern Tethys. 639–644. In SHERIDAN, R. E. et al. (eds). Initial Reports of the Deep Sea Drilling Project, 76, 947 pp. SALFELD, H. 1924. Die Bedeutung der Konservativsta¨mme fu¨r die Stammesentwicklung der Ammoniten. Max Weg, Leipzig, 16 pp. SCHOLZ, G. 1979. Die Ammoniten des Vracon (Oberalb, Dispar-Zone) des Bakony-Gebirges (Westungarn) und eine Revision der wichtigsten Vracon-Arten der west-mediterranen Faunenprovinz, Teil 1. Palaeontographica, Abtei- lung A, 165, 1–80. SHIMIZU, S. 1934. Ammonites. 1–82. In SHIMIZU, S. and OBATA, T. (eds). Cephalopoda. Koza chishitsugaku oyobi Koseibutsugaku, Tokyo, 137 pp. [In Japanese]. SHIPBOARD SCIENTIFIC PARTY 1998a. Site 1052. 241–302. In NORRIS, R. D. et al. (eds). Proceedings of the Ocean Drilling Program, Initial Reports, 171B, 1053 pp. —— 1998b. Introduction. 5–10. In NORRIS, R. D. et al. (eds). Proceedings of the Ocean Drilling Program, Initial Reports, 171B, 1053 pp. —— 1998c. Site 1050. 93–169. In NORRIS, R. D. et al. (eds). Proceedings of the Ocean Drilling Program, Initial Reports, 171B, 1053 pp. SPATH, L. F. 1921. On Cretaceous Cephalopoda from Zululand. Annals of the South African Museum, 12, 217–321. —— 1925a. Sur quelques ammonites du Gault, nomme´s par P. Reyne`s. Annales de la Muse´e d’Histoire de Naturelles Marseille, 20, 95–106. —— 1925b. On Upper Albian Ammonoidea from Portuguese East Africa, with an appendix on Upper Cretaceous ammonites from Maputoland. Annals of the Transvaal Museum, 11, 179–200. —— 1927. Revision of the Jurassic Cephalopoda fauna of Kachh (Cutch). Memoirs of the Geological Survey of India, Palaeontologia Indica, New Series, 11, 1–71. —— 1941. A monograph of the Ammonoidea of the Gault. Part 14. Monograph of the Palaeontographical Society, 94, 609–668, pls 65–72. SUESS, E. 1865. U¨ ber Ammoniten. Sitzungsberichte der Akademie der Wissenschaften Wien, Abteilung 1, 52,71–90þ 305–322. WESTERMANN, G. E. G. 1996. Ammonoid life and habit. 607–707. In LANDMAN, N. H., TANABE, K. and DAVIS, R. A. (eds). Ammonoid paleobiology. Topics in Geobiology, 13. Plenum Press, New York, 857 pp. WIEDMANN, J. 1964. Unterkreide-Ammoniten von Mallorca. Abhandlungen der Mathematisch-naturwissenschaftlichen Klasse der Akademie der Wissenschaften und der Literatur Mainz, 1966, 155–256. —— 1966a. Stammesgeschichte und System der posttriadischen Ammonoideen (Ein u¨berblick) 1. Teil. Neues Jahrbuch fu¨r Geologie und Pala¨ontologie, Abhandlungen, 125, 49–79. —— 1966b. Stammesgeschichte und System der posttriadischen Ammonoideen (Ein u¨berblick). 2. Teil. Neues Jahrbuch fu¨r Geologie und Pala¨ontologie, Abhandlungen, 127, 13–81. —— 1988. Plate tectonics, sea level changes, climate – and the relationship to ammonite evolution, provincialism, and mode of life. 737–765. In WIEDMANN, J. (ed.). Cephalopods – present and past. Schweizerbart, Stuttgart, 765 pp. —— and DIENI, I. 1968. Die Kreide Sardiniens und ihre Cephalopoden. Palaeontographia Italica, 64 (New Series 34), 1–171. —— and NEUGEBAUER, J. 1978. Lower Cretaceous ammonites from the South Atlantic Leg 40 (DSDP), their stratigraphic value and sedimentologic properties. 709–734. In BOLLI, H. M., RYAN, W. B. F. et al. (eds). Initial Reports of the Deep Sea Drilling Project, 40, 1002 pp. WRIGHT, C. W. 1996. Cretaceous Ammonoidea. 1–362. In KAESLER, R. L. (ed.). Treatise on invertebrate paleontology, Part L, Mollusca 4 (4). Geological Society of America, Boulder, and University of Kansas Press, Lawrence. —— and MATSUMOTO, T. 1954. Some doubtful Cretaceous ammonite genera from Japan and Saghalien. Memoirs of the Faculty of Science, Kyushu University, Geology Series, 4, 107–134. —— and KENNEDY, W. J. 1984. The Ammonoidea of the Lower Chalk, Part 1. Monograph of the Palaeontographical Society, 137, 1–126, pls 1–40. —— 1994. Evolutionary relationships among Stoliczkaiinae (Cretaceous ammonites) with an account of some species from the English Stoliczkaia dispar Zone. Cretaceous Research, 15, 547–582. —— 1995. The Ammonoidea of the Lower Chalk, Part 4. Monograph of the Palaeontographical Society, 149, 295– 319, pls 87–94. —— 1996. The Ammonoidea of the Lower Chalk, Part 5. Monograph of the Palaeontographical Society, 150, 320– 403, pls 95–124. YOUNG, K. 1979. Lower Cenomanian and late Albian (Cretaceous) ammonites, especially Lyelliceridae, of Texas and Mexico. Bulletin of the Texas Memorial Museum, 26,vþ 1–99. —— 1984. Cretaceous ammonites (Valanginian to Cenomanian) from Deep Sea Drilling Project cores, southeastern Gulf of Mexico. 709–734. In BUFFLER, R. T. et al. (eds). Initial Reports of the Deep Sea Drilling Project, 77, 747 pp. LEHMANN: UPPER ALBIAN AMMONITES 61

ZITTEL, K. A. von 1884. Handbuch der Pala¨ontologie. I. Abteilung Pala¨ozoologie. II. Lieferung. III. Cephalopoda. Oldenbourg, Munich and Leipzig, 329–522. —— 1895. Grundzu¨ge der Pala¨ontologie (Pala¨ozoologie). Oldenbourg, Munich and Leipzig, vii þ 972 pp.

JENS LEHMANN Institut und Museum fu¨r Geologie und Pala¨ontologie Universita¨tTu¨bingen Sigwartstrabe10 72076 Tu¨bingen, Germany e-mail [email protected]

Present address: Department of Geology University of California Typescript received 5 October 1998 Davis, CA 95616–8605, USA Revised typescript received 30 April 1999 e-mail [email protected]