The ammonoid fauna of the Kimmeridgian-Tithonian boundary beds of Mombasa, Kenya

ROYAL ONTARIO MUSEUM LIFE SCIENCES PUBLICATIONS INSTRUCTIONS TO ALTHORS

LIFE SCIENCES CONTRIBUTIONS R O Y A L O N T A R I O MUSEL'M

NUMBER 135

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IS

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HARISH M. VERMA G. E. G. WESTERMANN

The Ammonoid Fauna of the Kimmeridgian-Tithonian Boundary Beds of Mombasa, Kenya

ROYAL ONTARIO MUSEUM PUBLICATIONS IN LIFE SCIENCES

Contents

The Ro\al Ontano Museum publ~shesthree senes In the L ~ f eSclences LIFE YIE\CES CO\TRIBLTIO\S. a numbered serles of onglnal sc~ent~tic publlcat~onclncludlng monopaph~c uorLs LlFE SCIE\CES OCC4SIO\4L P4PERS. a numkred senes of onglnal sclentlfic publ~cat~ons prlmarll) shon and usuall? of taonomlc slgn~ficance LIFE SCIE\CES \IISCELL*\EOI s PLBLICATIO\S. an unnumbered serler of publ~catlonsof \ar~edsubject matter and format

Abstract 1 Introduction 2 Materials and Methods 4 Previous Work on Upper Jurassic of Mombasa Area 4 Upper Jurassic in Vicinity of Freretown 9 Previous Work 9 New Work 13 Age and Affinity of Freretown Ammonoid Fauna 17 lntroduction to Kimmeridgian and Tithonian Stages 17 Age of Freretown Fauna 18 Upper Assemblage 19 Lower Assemblage 22 Ecology and Biogeography of Freretown Ammonoid Fauna 23 Introduction 23 Ecology 24 Biogeography 25 Systematics 27 Suborder Phylloceratina Arkell, 1950 27 Family Phylloceratidae Zittel, 1884 27 Subfamily Phylloceratinae Zittel, 1884 27 Phylloceras isoypum Benecke, 1865 27 Subfamily Calliphylloceratinae Spath, 1927 28 Calliphylloceras benacense (Catullo, 1847) 28 C . aff. C . malqvanurn (Boehm, 1907) 29 Prychophylloceras ptychoicum (Quenstedt, 1845) 29 Holcophylloceras polyolcum mesolcum (Dietrich, 1925) 3 1 Suborder Lytoceratina Hyatt, 1889 32 Family Lytoceratidae Neumayr, 1875 32 Subfamily Lytoceratinae Neumayr, 1875 32 Lytoceras fraasi DacquC, 1910 32 Pterolytoceras montanum (Oppel, 1865) 33 Suborder Ammonitina Hyatt, 1889 34 Superfamily Haplocerataceae Zittel, 1884 34 Family Oppeliidae Douville, 1890 34 Subfamily Taramelliceratinae Spath, 1928 34 Taramelliceras ( T . ) cf. T. kachhense (Waagen, 1875) 34 T. ( T . ) nachynotum (Oppel, 1862)? 35 Subfamily Streblitinae Spath, 1925 36 Srreblites habyensis Spath, 1928 36 Superfamily Perisphinctaceae Steinmann, 1890 37 Family Perisphinctidae Steinmann, 1890 37 Subfamily Perisphinctinae Steinmann, 1890 37 Pachysphinctes beyrichi (Futterer, 1894) 39 Kutroliceras porringeri (Sowerby, 1840) 42 K . aff. K . potringeri 9 43 K.? cf. K . bathyplocus (Waagen, 1875) 3 44

All manuscrlpls considered for publicat~onare subject lo the scrutln) and editor~alpolicies of the Life Sciences Editorial Board. and to re\leu b) persons outside the Museum staff uho are authorit~esIn the panicular field invol\.ed. LlFE SCIENCES EDITORIAL BO4RD Senlor Edltor J R r+\cstrr Ed~tor D 4 L E R C4LDER Ednor JO\ BXRLOU External Ed~torc s CHLRCHER Manuscript Edltor RICHXRD u I\TERBOTTO\I Production Edltor 4\DRE4 G4LL4GHER ELLIS H \ R I S H \I \ E R \ I ~IS

a Research A~soc~ate of the Department of lnvenebrate Palaeontolog). Rojal Ontano

Museum G E G U+STER\l4V\ is Professor in the Depanment of Geology, McMaster University, Hamilton. Ontano. and a Research Associate of the Depanment of lnvenebrate Palaeontology, Royal Ontario Museum

Canadian Calaloguing in Publication Data Verma. Harish, M. Ammonoid fauna of the Kimmeridgian-Tithonian boundary beds of Mombasa, Kenya (Life sciences contributions, ISSN 0384-8159; no. 135) Bibliography: p. 116 ISBN 0-88854-297-6 I . Ammonoidea. 2. Paleontolog! - Jurassic. 3 Paleontolog! Kenya - Frere Tonn. I. Westermann, G. E. G. (Gerd Ernst Gerald). 1927- 11. Royal Ontario Museum. 111. Tltle. IV. Series.

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QE807.A5V47

564.53'0967623

C83-098584-0

Publ~cationDate: 28 March 1984 ISBN 0-88854-297-6 ISSN 0384-8159 @The Rolal Ontano Museum. 1984 100 Queen's Park. Toronto. Canada M5S 2C6 PRI\TED 4 \ D BOL\D I \ C 4 \ 4 D 4 47 THE 4LCER PRESS

Subdichotomoceras(?) cf. S. sparsiplicatum (Waagen, 1875) 46 S.? aff. S. sparsiplicatum (Waagen, 1875) 47 Family Ataxioceratidae Buckman, 1921 48 Lirhacoceras cf. L. fraasi (Dacqui, 19 10) 48 L. ? mombassanum (Dacqut, 1910) 49 L. cf. L. olbulum (Qwnstedt, 1887) 49 Phanerosrephanus (Nothosrephanus) digoi sp. nov. Q 53 Family Aulacostephanidae Spath, 1924 57 Procraspedites cf. P. africanus (Zwierzicky, 1914) Q 58 Grai,esia loupetinei sp. nov.9 59 G. aff. G. loupekinei sp. nov.9 63 Family Aspidoceratidae Zittel, 1895 63 Subfamily Aspidoceratinae Zittel, 1895 63 Aspidoceras cf. A. acanrhicum (Oppel, 1863) 9 + ?d 63 A. aff. A. acanthicum (Oppel, 1863) 66 A. iphiceroides Waagen, 1875 9 + 8 66 A. cf. A . iphicerum (Oppel, 1863) 68 A . cf. A. apentlinicum Zittel, (1869) 1870 69 A. (Ph.vsodoceras) cf. A. avellanum Zittel, 1870 70 A . (P.) circumspinosum (Quenstedt, 1858) 71 Family Simoceratidae Spath, 1924 72 H~bonoricerashybonotum (Oppel, 1863) 72 H. cf. H. hybonorum (Oppel, 1863) 73 H . cf. H . ornatum (Spath, 1931) 73 H. pressulum (Neumayr, 1873) 75 H. cf. H. ciliarum Berckhemer & Holder, 1959 76 Acknowledgements 77 Plates 78 Literature Cited 116

The Ammonoid Fauna of the Kimmeridgian-Tithonian Boundary Beds of Mombasa, Kenya Abstract The macroinvertebrate fauna of the poorly to moderately exposed upper Changamwe Shales at Freretown, north of Mombasa, was collected with the best precision possible. The sequence consists of 370 to 600 m of silty shales with layers of calcareous concretions and lenticular siltstone. These have yielded a rich, almost exclusively ammonoid fauna in parts of the upper half of the section and a scarce to moderately rich similar fauna through the remainder of the section. This collection of 325 specimens was placed in 28 species, 2 of them new. The rich and diverse upper assemblage consists of species of Phylloceratina (c. 20%) Phylloceras, Calliph.vlloceras, Pt.vchophy1loceras, Holcophylloceras, Lytoceratina (c. 5%) Lyroceras. Prerolytoceras; and Ammonitina (c. 75%) Taramelliceras, Srreblires, Katroliceras, Pachysphincres, Subdichotomoceras?, Lirhacoceras, Aspidoceras (s. srr.), A. (Physodoceras), and significantly the first findings in Africa of the genera Procraspedites, Phanerosrephanus with P . (Norhostephanus) digoi sp. nov., Gravesia with G. loupekinei sp. nov., and of the species Hybonoticeras hybonotum. The association of Gravesia and H. hybonotum clearly identifies this assemblage as belonging to the Hybonotum Standard Zone ("Gravesia Zone") of the basal Tithonian, hitherto restricted to the Mediterranean Province (Tethyan Realm). This is the first good evidence of both zonal indices and thus of the Hybonotum Zone for the Indo-Southwest Pacific Province; it clearly dates the coeval Madagascan "Zone of H . hybonorum and Aspidoceras acanthicum" and the "2nd Katrol Assemblage" of Kachchh, India. Significantly, this "Coroa Mombasa assemblage" was previously dated at latest Kimmeridgian (s. srr.), about Beckeri Zone. Faunistic comparison of this assemblage within the Ethiopian Subprovince, Indo-Southwest Pacific Province, indicates a somewhat closer affinity to Madagascar than to Kachchh, and a closer connection than previously assumed of (only) the East African-Madagascan faunas with the Mediterranean Province, in particular, the Submediterranean Subprovince of Northwest Europe. Gravesia is well known from as far north as the Northwest European Subboreal Province, and probably the northern Ural Mountains of the Boreal Realm. The lower assemblage consists of Kimmeridgian-basal Tithonian taxa common in the Indo-South~estPacific Province: Phylloceratidae, Taramelliceras, Katroliceras, Pachysphincres, Lirhacoceras, and

.4spidoceras (s. str. ), with several species identical to those in the upper assemblage, as well as H?bonoticerus pressu!um. an index of the (lower) Beckeri Standard Zone in (Sub-) Mediterranean Europe. These beds are probably contemporaneous with the Madagascan "Zone ofAspidoceras longispinurn and ulterneplicutus" and with the "1st Katrol Assemblage" of Kachchh. The other similar northeast African faunas need reinvestigation. The basal Tithonian is probably disconformabl> overlain b) the mid-Neocomian Freretown Limestone Group. with the hiatus spanning most of the Tithonian and the basal Cretaceous.

Introduction Recent advances in our understanding of the earth's crustal behaviour have resulted in the rejuvenation of the "continental drift" hypothesis. It is therefore desirable to take a fresh look at the palaeobiogeographic data and their compatibilit) with the pre-drift reconstructions. Some of the areas greatly affected by the early Mesozoic break-up of Gondwanaland are the east coast of Africa, Madagascar, peninsular India, Antarctica, and Australasia. Our knowledge of the Jurassic biostratigraphy and palaeobiogeography of these areas is, however, far from perfect and, for the eastern part of the Jurassic Tethys, dates largely from the late 19th and early 20th centuries. Important beginnings toward a revision of the late Jurassic faunas in some of these areas have been made in the last three decades, beginning with Arkell's (1956) grand synthesis of the World Jurassic. For instance, the fauna of the Spiti Shales in the Himalayas, previously known only from the works of Uhlig (1903, 1910), was recently in part re-collected and described by Helmstaedt (1969) and Mouterde (1971). Some of the famous Jurassic faunas of Kachchh described by Spath (1927-33) were re-examined by Agrawal (1956, 1957). Contributions by Stevens (1963a, 1963b. 1965, 1967, 1968, 1971), Westermann and Getty (1970), and others (see references in Sato, 1975) have advanced our understanding of the palaeobiogeography and biostratigraphy of the Australasian region. The Jurassic of Madagascar has been the subject of several monographic works by Besairie and Collignon (1956), Collignon (1959, 1960, 1962), and Besairie ( 1936, 1972). while the Ethiopian fauna was investigated by Zeiss (1971, 1974). The ammonite-bearing intervals of the Tendaguru Group of Tanzania were studied by Zwierzicky (1914) and Dietrich (1925, 1933) and are in need of revision. Dr. M.K. Howarth (pers. comm.) of the British Museum has re-examined the known ammonite occurrences of Tanzania and will revise the faunas. It was therefore considered highly appropriate to reinvestigate the Jurassic ammonite faunas of Kenya hitherto known mainly through the works of Beyrich (1877, 1878). Futterer (1894), Dacque (1910), Gregory (1921), and Spath (1930a). and MacKinnon Wood (1930, 1938) in the "McKinnon Wood" monographs. One of us (Westermann) visited coastal Kenya during his sabbatical leave in 1973 with a view to reinvestigating the Middle and Upper Jurassic biostratigraphy and subsequently reported on the Bajocian ammonoid fauna (Westemann, 1975). In the present paper we describe some important new faunal elements from the Upper

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L Text-fig. 1 Locality map and geological map of the Mombasa area. The stud? area lies within the heav) rectangle 1 to 3 km north of Freretoun.

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lenticular interbeds of grey limestone and siltstone. The lower pan of the formation bears the Indo-Southwest Pacific Mayaitinae fauna, with Perisphinctes (s. I . ) , Ma\.a;te.$ (including Epima~aites).Dhosoites, and Progre~.icerus,all of which Arkell (1956:322-23) dated as Transversarium Standard Zone [ Z Plicatilis Zone]. Middle Oxfordian. However, the Lower Oxfordian Cordatum Chronozone seems also to be present at an undetermined level as indicated b! the single Puro~~rdekit~diu sp. and Eltaspidoceras cf. acuticosratum (Young and Bird) (Arkell. 1956). This fauna has also been re-collected but is not being revised here since studies of the similar Indonesian fauna have been initiated by one of us (Westermann et al., 1978). The late Kimmeridgian to basal Tithonian assemblage of the middle and upper parts of the Changamwe Shales is discussed below in detail.

Upper Jurassic in Vicinity of Freretown PREVIOUS WORK Since Cretaceous outcrop is exceedingly rare in the marine embayment of Kenya, Haw (1964) reinvestigated the stratigraphy of the Freretown area, especially in the light of the microfaunas. According to Haw's unpublished report and the accompanying maps (partly reproduced here with modification. by permission of the British Petroleum Company; Text-figs. 2, 3), the following stratigraphic units are exposed in the Freretown-Mto Panga area:

< 30

Shales, massive calcareous grey with occasional silty beds, sandstone and layers of hard calcareous mudstone to argillaceous limestone. Rich planktonic microfauna and benthic coral-bivalve fauna. Albian.

(5 m)

Sandstone, hard, calcareous, fine-grained

25 m

Mudstone, soft, clayey, calcareous. - (Upper) Aptian.

Freretown Limestone Group, Freretown Limestone s.s.: 8m Detrital limestone, very finely calcarenitic, bioclastic, some interclastic sparite, generally well bedded but many bedding planes irregular (minor diastems); at top with silty interbeds. Coral-bivalve assemblage mainly near top; limestones with Orbitolinoides hulgarica janeshci. - (Barremian? - Lower) Aptian. Freretown Limestone Group, lower part: Silty, sparry limestone. Abundant macrofossils: E x o g ~ r o minos, Osrrea , corals. - "Neocomian".

( 5 m)

Changamwe 30 m (60 m according to Haw, fig. 4)

Shales: Massive dark shales, slightly calcareous with occasional calcareous partings (concretions absent). With rare ammonites, that is, "Groebericeras" [ = Procraspedires, Lower Tithonian] "?Preroi~toceras" [?Tith.-Valang.]; microfauna contains "Cretaceous Marssonella and G~roidinain an otherwise Kimmeridgian shale-type assemblage." -

"Neocomian". Sample No. 1781 with the supposed Berriasian "Groebericeras", from about the lower third of the section, was. however, said to have a Kimmeridgian microfauna, while the on11 higher Sample No. 1790 in the middle of the "Neocomian" interval yielded a rather poor microfossil assemblage dated as "Kimmeridgian-Neocomid . (250 m?)

Massive shales. dark gray with greenish blue (fresh), generall) slightly calcareous, and with occasional very impersistent muddy limestones with very calcareous shales and fairly common lime! concretions and septarian nodules. Frequently containing ammonites. - "Lower Kimmeridgian-Tithonian".

Remarks Haw (1964) placed the upper 401120 m of this shale series (40 m according to fig. 4; 120 m according to text and fig. 3) in the "Middle-Upper Kimmeridgian" and "Tithonian". However, the critical "Tithonian" Sample No. 1760 contained only a "Phwodoceras 1" identified and dated "Kimmeridgian" by Eames and Clarke (1964). while Sample No. 1780 of the supposed "Upper Kimmeridgian" included the same ammonite species together with a rich microfauna identified and dated as "Kimmeridgian" (Eames and Clarke, 1964). The dating of the entire fauna was actually based on the associated ammonites, since the microfauna "is not distinguishable from the Lower Neocomian samples apart from the absence of Cretaceous genera and species" (Eames and Clarke, 1964). The boundary LowerIMiddle Kimmeridgian (s. angl.) was placed according to ammonite evidence, that is, the assumed restriction of Aspidoceras iphiceroides and Lithacoceras kenpense to the Lower Kimmeridgian (s. angl.). The approximate boundary was inferred to run with general northwest-southeast direction through the New Mto Panga Quarry. This coincides approximately with our KimmeridgianTithonian boundary. We have found another specimen of Procraspedites ["Groebericeras"] in an assemblage of the Hybonotum Zone, basal Tithonian. Since the supposed ammonite evidence for a Neocomian age of the uppermost Changamwe Shales does not exist, the evidence rests on the remaining fauna. The (other) megafauna consists of long-ranging corals and bivalves and aHibolires, to which Eames and Clarke (1964) attribute "Neocomian" (I sp.), "Neocomian-Aptian" (3 spp.), and "Lower Cretaceous" (7 spp.) ages. The microfauna was said to indicate mostly Albian age; but the lowermost Sample No. 1778 came from the lower Freretown Limestone which was dated "Neocomian". Evidence for a "Neocomian" age of the shales subjacent to the Freretown Limestone was, therefore, entirely dependent on the microfauna, contrary to the recent statement by Walters and Linton (1973: 140). The "Lower Neocomian" date given by Eames and Clarke (1964) rests on the association of long-ranging Kimmeridgian-Neocomian and Cretaceous foraminifera species, for example, Ammodiscus, Gaudr~ina,G~roidina, Marssoriella, Barhysiphon, Lenticulina, and his "Lower", rather than "Upper", Neocomian date was apparently based on the supposed occurrence of Groebericeras rather than on his own work, The stratigraphic interval between the base of the Freretown Limestone and the supposedly Lower Neocomian samples is, furthermore, unknown. The limestones

o,tcmp only in the small Freretown quarr). and the thickness of the uppermost Changamwe Shales has probably been overestimated previously. being based on an 10- to 15-degree regional southwest dip. The neu trench for the water pipeline (Text-fig. 2) directly north of the Cretaceous exposures shou s horizontality. w e suggest therefore that the earl) Cretaceous samples in\estigated by Eames 1 ) came from immediatel) subjacent, if not from the base of the F r e r e t o ~ nL~mestone and 2) belong in the Upper, rather than Lower. Neocomian Consequentl>. the most probable position of the disconformity remains near the base of the Freretoun Limestone Group, with the hiatus spanning "Middle" Tithonian to Lower Neocomian.

NEW WORK In the vicinity of the New Mto Panga Quarrq. the flat-l) ing Changamwe Shale forms a gently undulating surface dissected b) a few shallow valleys. Although the $hales are mostly covered with talus and allilvium, they are exposed in the quarry. Several bands of small to large calcareous concretions, however. crop out frequently. often forming the tops of small hills or ledges along gentle grass-covered slopes. They are best exposed and fossils most easily found where the pastures have been freshly burned. The clay quarry, however, is operated bl surface scraping, since its depth is limited by the high groundwater level. It is several hundred metres in subcircular diameter, but up to only several metres deep and the bedding structures of the soft, weathered shale have been obliterated. The only possible useful markers to determine attitudes are therefore the layers of concretions and lenticular limestones. Fortunately, a trench 1 to 1.5 m deep had been excavated for a water pipeline along the western edge of our area, exposing the flat-lying shales with concretions continuously for more than a kilometre (Text-fig. 2). The total section, therefore, can only be reconstructed from disjunct outcrops of concretionary layers, and some mostly small shale exposures. Consequently, the measurement of stratigraphic thickness is subject to considerable error. In view of this, we estimate the thickness of the Changamwe Shale of the Freretown area to be somewhere between 220 m and 400 m. At least another 100 m of Kimmeridgian shales and several hundred metres of Oxfordian shales are present around Mombasa. Although the attitudes are rather persistent, with horizontality in the southwest (and ?west where the trench is parallel to supposed strike) and a dip of 10 to 15 degrees west-southwest around the quarry and probably to the north (also the Freretown Limestone in the south), faults may remain undetected under the poor outcrop conditions. Fortunately, the main traverse is from south-southwest to north-northeast, which is parallel to the major normal fault system of the Mombasa area, for example, the Freretown Fault (Caswell, 1956, fig. 14) and the "Hypothetical fault" of Haw (1964, figs. 1, 2). The smaller east-west fault between the "Upper Jurassic Shales" and the Freretown Limestone Group (Linton and Maclean. 1955, fig. 2: Haw. 1964) also does not affect our stratigraphic section (locs. 8a-8k) Localities 8a-8k: Freretown Section (Text-fig. 2, Table 2)

Vicinity of New Mto Panga Quan) of Bamburi Portland Cement Works. I to 3 km north of Freretown, Mombasa area (Text-fig. I). The locality descriptions are

arranged stratigraphicall) (top to bottom) and the nonexposed intervals are indicated in parentheses (Text-figs. 2. 3). The species are listed in Table 2. Freretown Limestone: M'orL For description see under Pre~~ioris Changamwe Shales: (10 to 150 m) 8f-8g: Grejish shales with some large calcareous concretions, exposed in 5 to 8 m pipeline trench: attitude horizontal (at least at bend of trench) to near-horizontal. Holcopli~llocerus.Procraspedires, H~bonoticerascf. It~bo~~orron. &: 0.5 to 1 m

8d : 0.5 to 1 m

(30 to 50 m shales) La!.er of calcareous concretions and lenticular siltstone forming flat ridge; probably with sentle southwest dip. Fossiliferous: abundant Holropl~~llocerus,Phanerosrepl~anus (Nothostephanus) digoi , and Aspidocerus ( P h ~ s o d o c e r a s ) , also Gra~.esio loupekinei and Sot11a1i11uurilu.i. (3 to 5 m "shales") Layer of siltstone and concretions, exposed on gentle slope; attitude near-horizontal. Highly fossiliferous: abundant Phylloceratidae, Aspidoceras (s. str. ). Srreblires, and Pachysphincres , also Lytoceratidae, S~rbdichoromoceras?,A. (Pl~ysodoceros),Gravesia loupekinei, Tarantelliceras, and Hybo~~oriceras cf. hybonorum.

Shales with concretions, poorl) exposed on the north side of same fossiliferous: Pt~cltoph~llocera.~, Tarcr~rlelliceras. Katro1icera~-?,Paclt~.\ph~nctes, H~hortotic.erassp

8j: ,, 10 to I5 m hillock as 8h. Moderatell

8)i:

c. 5 m

Total about 300 to 350 m, closely approximating the earlier estimates for Kimmer~dgian(s. angl. ).

Locality 15 (Text-fig. 1) Changamwe Peninsula, Kipevu, about 400-m-broad cut slope behind the new docking facilities west of the Kipevu Causeway in the northeast comer of Port Reitz. Att~tudemostly near-horizontal and dipping east at west end of outcrop. The section is from top. c. 7 m

Thick lenticular beds of sandstone and siltstone with shale) interbeds and sandstone dykes; unfossiliferous.

8m

Shale; unfossiliferous

3.5 m

Shale with sandy concretions and ?sedimentary dykes; unfossiliferous.

15: 10 m

Blue-grey friable shale; near top moderately fossiliferous: P n c h o p h ~ l loceras prychoicum, Lyroceras fraasr. Prerolvroceras morltunum. Perisphinctinae indet . , Aspidoceras (Aspidoceras) cf. hoplisum. A . (Physodoceras) cf. avellanum, Taratnelliceras cf. kachhense.

15b: c. 10 m

Shale with fossiliferous concretions, ferrugineous partings. Pterolytoceras pychoicum, Lyroceras fruasi. Aspidoceras (Pseudou.augenia) aff. haynaldi, Perisphinctidae indet., Belemnopsis ranganettsis (Futt.).

15a: C. 5 m

Shale with concretions; moderately fossiliferous: ?Pach~sphinctes beyrlchi, Srreblires habyensis.

C.

80 to l I0 m Greenish and greyish shales with some concretions. Poorly exposed northwest of quarry; lower part in west side of quarry. &:

Concretions in upper half with: Calliphylloceras, Srreblires, Lithacoceras, Pach~sphincres?,Karroliceras, Subdichoromoceras?, Aspidoceras, A. (Ph?sodoceras), and Hybonoriceras hybonotum.

8a (=81=4): Near base moderately abundant: Phylloceratidae, Pterolyroceras, Srreblires, Pachysphincres, Lithacoceras?, Aspidoceras (s. srr.), and H~borloricerashybonotum. 8b: 5 to 10 m

Poorly exposed, mostl) shales with concretions; at top (8bl) irregular lenticular limestone bed and large concretions, dipping 12-15" WSW. Concretions and limestone moderately fossiliferous: Phylloceratidae, abundant Aspidoceras (s. s n . ) , also Karroliceras. Pachysphincres. Subdichotomoceras?, Lirhacoceras, Hybonoriceras pressulum , and Paracenoceras. (40 to 70 m "shales")

8h: c . 10 m

Shales with bands of concretions, slightly exposed on gentle south slope of hillock several hundred metres north of the bend in the road leading around the quarn; probably with gentle southwest dip. Moderately fossiliferous: Calliphylloceras, Taramelliceras, Pachysphincres?, Karroliceras, Lithacoceras?, Hybonoriceras pressulum , and abundant Aspidoceras (s. srr.).

(''50 to 100 m) Calcareous shales with some concretions in stream cut approximatel) 0.6 hm northeast of 8g; near-horizontal. Scarce fauna: .4spidoceras (s. srr. ), Lirhacoceras?

Total about 43.5 m.

Localit! 7 Changamwe Peninsula, Kipevu, section cut b) main road (from traffic circle) between ridge over railroad and dock gate, 0.2 to 0.5 km northwest of locality 15. c. 10 m

Shales with concretions, topped b) some sandstone and siltstones. Probably same beds as included at localit) 15. Moderately fossiliferous: Aspidoceras (Pseudo\r~aagenia) aff. haynaldi (Neumayr), Lirhucoceras cf. arussiorum (Dacque), L. cf. sigali (Collignon), Taramelliceras trachxnorum (Oppel)?, Perisphinctidae, ? P a c h ~ sphincres fragments.

Age arzd ASJinir~of Frereto\c+n Amlnorloid Fauna INTRODL'CTION T O KIMMERIDGIAN .4ND TITHONlAN STAGES The Stages Oxfordian. Kimmeridgian. and Ponlandian as defined b! d'Orbignj ,1842-51) were named on the basis of the faunal elements existent in northwest Europe and can be universally recognized and correlated only up to the top of the ~immeridgian(s. srr.) (Crussolian) Beckeri Zone. The post-Kimmeridgian (s. srr. or and A . angl.) stages have been called Portlandian and Volgian in the Boreal Realm and mostly, though not always, Tithonian over the rest of the world (Table 3). Disagreements have revolved around the time-stratigraphic and geographic definitions of the stages and the time equivalence of their upper and lower limits (see Verma and Westermann, 1973: 159, table 7). Confusion has arisen where. for want of agreement over which name should be used for which realm, the standard English stage names have been used for areas belonging to the Tethyan Realm. Some uniformity and international agreement over stage usage have been achieved as a result of three successive colloquia held in Luxembourg in 1962 (Zeiss, 1962, 1964; Maubeuge, 1964) and 1967 (Zeiss, 1971) and in Budapest In 1969: Kimmeridpian and Portlandian are to be used as originally defined in southern England for areas of northwest Europe, with the boundary between the Rotunda (nor Pallasioides) and Albani Standard zones; Tithonian should be used for all areas of Tethys including Submediterranean Europe and all (other) Perigondwanian areas including those formerly ascribed to the Pacific Realm (i.e., Japan, Mexico, Cuba, southern Andes, Antarctica, New Zealand, etc.). The Tithonian faunas are extremely extensive and recognizable over a larger part of the globe than those of Upper Kimmeridgian (s. an$. ), Portlandian, and Volgian. They offer, therefore, much greater possibilities of correlation (Enay, 1963:6, 9; 1964, tables 1-4; 1973; Zeiss, 1968, 1977b. 1979, 1982; Oloriz and Tavera, 1981). The lower limit of the Tithonian Stage, defined by the base of the Hybonotum Standard Zone, is marked by the appearance ofH~honorirerash~bonorurn(Oppel) or, for want of this species, by the genus Grarvsia, particularly the species G . gra~esiuna(d'orbigny) (Enay, 1963, 1964, 1972, 1973; Barthel and Geyssant, 1973; Ens) and Geyssant, 1975). The upper limit is somewhat more difficult to define and complicated by the fact that it also marks the boundary between the Jurassic and Cretaceous systems. The Volgian corresponds in time to the Tithonian and the zonal correlation is shown by Zeiss (1968, tables 5, 6; 1974; 1983). In the entire Tethyan and Perigondwanian Realms, therefore, most of the "Middle Kimmeridgian (s. )" (post-Beckeri Zone) should be included in the Lower Tithonian (s.1.). In the Tethyan and Volgian Realms, the "Lower Kimmeridgian s. ongl. " thus becomes the Kimmeridgian (s. srr.) (Ziegler, 1977). or simply, Kimmeridgian as used here. The use of the substage name Crussolian for this unit was also proposed at the first Luxembourg colloquium in 1962 (Zeiss, 1962, 1964). but has found little acceptance, tending to confuse the matter further. The Tithonian can be divided into two (Spath, 1950; Arkell, 1956: Zeiss, 1968; Verma and Westermann, 1973; Enay and Geyssant, 1975; Wiedmann, 1980; etc.) or three substages (Donze and Enay, 1961; Enay, 1964, 1972, 1973; Zeiss. 1983). The bipartite subdivision is favoured by us on account of the difficulty of recognizing a distinct "Middle Tithonian" assemblage (Zone of Pseudolissoceras zirreli in Mexico, >

i B EZONE CKERI i

I

HYBONOTUM

ZONE

- K I M M E R I D G I A N ~ ~T~l T~ ,H O N l A N t

(Crussolionl

I

j I .

i

Table 2 The stratigraphic occurrence and ranges of the various species of ammonites within the Kimmeridgian-Tithonian boundaq beds of the Mombasa area. The thickness of the lines indicates relative abundance. Dots denote single occurrence. Dashed lines indicate presumed occurrence. Letters a to I, refer to localities indicated in Text-fig. 2 .

EUROPE (BOREAL)

I

(TETHYAN)

. . , central and southern European after Enay and ~ e ~ s s a (1975), nt Zeiss (1977b, 1979, 1983), and Oloriz and Tavera (1981); Madagascar after Collignon (1964). Argentina, and Iraq, Zone ofSemiformiceras semiforme in the Alps) in all parts of the globe (Verma and Westermann, 1973, table 4; Enay, 1973, table 1). The recognition of a "Middle" Tithonian assemblage in northern Iraq by Spath (1950) on the basis of Pseudolissoceras d e l i in association with his Phanerostephanus-Nothostephnus fauna, conflicts with our discovery of Hybonoriceras hybonorum and Gravesio loupekinei sp. nov. together with similar faunas from Kenya. As pointed out by Enay (1972), only a bipartite classification is acceptable when utilizing the common elements of correlation; that is, Hildoglochiceras is known from the Himalayas, Madagascar, Tanzania, Cuba, and Mexico; Kossmario from the Himalayas, Indonesia, Australia, New Zealand, Argentina, Mexico, Texas, and California; and Lvrohoplires from Salt Range, Madagascar, Argentina, and Cuba. AGE O F FRERETOWN FAUNA The 220-to-400-m Changamwe Shale at Freretown bears a moderately abundant to highly abundant ammonoid fauna with overlapping vertical ranges in what appears to

be a rapidly and continuously deposited ner~tic5eries (Table I ). The stratigraphic of the ammonoid tala is. howeter. mostl! rporad~cra~herthan cc>ntinuous. ,ith most fossils occurring in a dozen or so horizons of concretions or lenticular lime\tone\ and siltstones. This I S particularl) true of the Inuer half and near the top of ,he sequence. Furthermore, the vertical ranees are rearunabl! uell Lnoun for fewer than 10 of the 34 recorded taxa. mostl) from the hlfhl! ios\iliferous middle and upper parts of the sections (Table 2 ) . The ammonoid fauna is relativel) uniform throughout the sequence. uith several species ranging throughout, for example. the Ph\lloceratidae Colliph~llo~~rrns he~luc.c*n.se (Catulio) and Pr~choph\~llocc~ri~s pr\c.lloirrrrrl (Quenstedt). as uell as =vzral Ammonitina, for example. Knrri~lic.crti.\porrirrgeri (Sou .). and the ahundant A.cprrloc.ef'il5rphireroides Waagen. Man! other taxa are represented b) single or a feu, specimens so that their range is unknown (Table 21. Both a lower and an upper ammonite assemblaee can. nevertheless. be clearl! distinguished. with a possible narrow o\erlap in bed 8b. The upper assemblage is b! far the richer in number of specimens and taxa

Upper assemblage with Graresia, Phanerosrephanus, and Hybonoticeras hpbonohrm This assemblage is characterized b) abundant Srrchlircs hobyc~risirSpath. Phcrtrerostephanus (Sorhostephurius) digoi sp. nov.. .4.\pidoc.~r-u.\(il~pidocer~rs) cf. (rcc111rhicurn (Oppel), and the less abundant Srthdic-horor~locer-ci.~? cf. .spar-siplicc~frrtn (Waagen). Aspidoceras (Ph~sodocero.c) cf. a~~ellcrri~o?~ Zittel. A . ( P .J circrrr~l.~pirlo.srrrn (Quenstedt). Gru~tesialoupekiriei sp. nov.. Procruspc~lilcs cf. ufriccr~rus(Zwierzicky). H~honoricerosh~bonotum(Oppel). and H . cf. ortlurrirll (Spath). The fauna also includes taxa of questionable ape sisnificance, such as Phylloceratina, in particular the rather ahundant Holcopllylloceras polyolcron (Benecke). and the rather scarce Lytoceratina: as well as the rare Tcrrarr7elliceras cf. kachherlse Spath; Kurrolireras porritlgeri (Sow.) and K . katrolertse (Waagen), of which at least the former ranges downward: Lirhococeras cf. ficrasi (Dacque): and Aspidocems iphiceroides Waagen, which is abundant in both assemblages. The presence of the two index fossils Hyhorloticerus h~botlorurnand Gravesia securel) places this assemblage in the Hybonotum Standard Zone or Gravesiu Zone. both of which have previously been essentially restricted to Europe (Zeiss. 1964, 1965, 1977: Enay, 1972. 1973: Sapunov. 1976. 1977: Barthel and Schairer, 1978). H . hxbonontm in beds 8a and 8c, and H . cf. /7\bonoru1n together with Gro~,esiain bed 8d and alone in beds 8f and 8g at the top of the sequence, estahlish the range of the zone. Even if Gruvesia is represented by the neu species G . Ioupekir~ei.the entire, distinct genus is restricted to the Hqbonotum Zone throughout its geographic range in Europe. and all previously known species are said to be penecontemporaneous (Barthel. 1959: Ziegler, 1960; Hahn, 1963: Cope and Zeiss, 1964: Ena!. 1966, 1973). Gra1.esia has also been recorded from the Russian platform and western Siberia where it marks the base of the Yolgian stage (Zeiss. pers. comm.) The apparent restriction of Gravesia to the narrou strar~graphicinterkal of beds 8d to 8e may be owing to its relative scarcit). Zeiss (19681 supposed, however. that Gruvesia is facies-controlled by shallow neritic to sublittoral limestone: the high]! fossiliferous limestone beds of the Freretown sequence are probabl) an example. The earliest occurrence ofH. hybor~orum(bed 8a) is superjacent to the latest record from bed 8b of

H. pressrtlro~t of the (lower) Beckeri Standard Zone (see below). The zonal (and stage) boundary is therefore between beds 8a and 8b. H~bor~oriceras ortlarlim Spark probably also from bed 8a, occurs in Kachchh, India. in both H\.honorirerus Zon (Spaih, 1931). The questionable H. ciliarlcm Berchhemer and Holder (1959) from bc . 8c was originall) described from the Beckeri Zone of German?, but the specie3 appears to be known from the holotype and one fragment only and thus is of unknown vertical range. The upper range of H\bo~roticeras in Europe coincides with that of Grovesia, that is, the upper boundary of the Hybonotum Zone (Zeiss, 1975). Of the other taxa apparently characteristic of the upper assemblage, Strebfiiez kabyr~sisoccurs also in the Hybonotum Zone of Madagascar (Collignon, 1959). b~ mainly in the Beckeri Zone of Kachchh (Spath, 1931). Subdichoromoreras? ct sparsipliea~ri~~r (U'aagen) occurs also with identical forms in the Tendaguru Group o; Tanzania (Zu-ierzicky, 1914, pi. 8, figs. 1, 2); but its stratigraphic level ("Nerineenschicht.') is uncertain since it was found by natives in a river bed supposedly maid}- in the "Nerineenschicht" (Janensch and Hennig. 1914:5). Dietrich (1933:20) lists "Perisphirzcres sparsiplicatus" from both the Smeei and Nerineen [.2'erit~ellulBeds. His small. figured specimen (pl. 1, fig. 2), however. resembles more closefr the dubious nucleus "Perisphittctes larissimus" [Path?sphi~lcres?]Zwierzicky (1914, pl. 8, fig. 4). found loose, while Dietrich's larger s w i m e n 11933. fig. 4) with similar coarse ribbing comes from the proximity of the Nerineila Bed. Similar poorly preserved specimens were illustrated from the Hybonotum Zone of Madagascar (Collignon, 1959, figs. 4, 7, 8). Pha~~erosrephat~us (Nothostephanus) digoi sp, nov., known only from bed 8e where it occurs in abundance, can be closely compared only with P. (A'.) kurdistarre~~sis (Spath) and, perhaps, P. (P.) subsene* Spath from a black limestone sequence ("bed i" = 10 m) in Kurdistan, Iraq (Spath, 1950). Although Spath placed that assemblage tenatively in the '.Middlem [upper Lower] Tithonian, he also reported a Hrbonoticeras ex gr. hybonotum of the same preservation which was reported to him to have come from the same interval. We conclude that the 10-m sequence of black limestone at Jebel Gara probably contains more than one zone, including the Hybonotum Zone. Aspidoceras cf. acanthicum, if identical with that species, appears to differ significantl) in age between western Europe and Kenya-Madagascar. The species has been regarded as an index for the Kimmeridgian "Mutabilis-Acanthicum Zone" of eastern France (cf. Contini and Hantzpergue, 1975) and of the Acanthicum Zone in southern Germany (Ziegler, 19771, preceding the Eudoxus and Beckeri Zones (Table 3). In Ken?a and Madagascar, this species is abundant in the Hybonotum Zone (Zone ofH. hxbonorum andA. acanthicum of Collignon, 1959). In Kachchh, India, it is rare in the ? Beckeri Zone (Spath, 1931), and probably ranges downward into that zone also in Kenya (Spath, 1930a). The similar A. longispinum (Sow.), occurs in the Eudoxus Zone (Contini and Hantzpegue, 1975) and may be conspecific with the poorly defined A. acanrhicum (cf. under that species). According to Wiedmann (19801, A . cf. aca~rthicumoccurs also in the Lower Tithonian of Peru. Aspidoceras fPh~sodoceras) avellanum occurs also in the Lower Tithonian of Mr. Catria in Italy (Zittel, 1870) and in the Hqbonotum Zone of Spain (Enay and Geyssant, 1975). It closely resembles A. (P.) pinini (Oppel) from the Solnhofen Limestone, southern Germany, in rhe same zone. The holotype of A . (P.) circurnspi~rosum, however, comes from the Lower Kimmeridgian of Germany (Maim gamma).

Karro)jceras (s. srr.) occurs in Submediterranean Europ: as earl) as Eudoxus Zone iasoutheast France (Contini and Hantzpergue. 1975). but ranges into the lower Hybnotum Zone in southern Franconia. German! (Zeiss. 1975). The similar subeenus ~Crussoliceras. however, which has k e n separated gnericall\ onl! ncentlj b) some authors (e.g., Contini and Hantzprgue. 1975: Sapuno\.. 1977: but not ziepler. 19771, marks already the Di\isun~ and Acanthicum Zones of southeastern France to Bulgaria. where Karroiicerus is str. ) appears to be absent t~apuno,,,1977: Contini and Hantzpergue. 19751. In Xladagzscar and Kachchh. on hand, Karrolicera~(s. J r r . ) appears to be restricted to the Hybonotum Zone ( a n e of ~ ~ ~ i d o c eacatlrhicrtrn ras and H. I~!bnrrotu~~~ of Collignon. 1959. and "2nd ~~~~l ,4s5emblage" of Middle Katrol Group of Spath. 1931). The typical k'. 6 . 1 L ~ ~ found ~ at~ Mombasa / ~ is also ~ quite ~ J abundant ~ in Madagascar and Kachchh and sjmilarly ahaociated with Parh\sphinrres . Questionable Kurroiiceras u ere also p-poned from the ' ' 1st Katrol Assemblage", basal Katrol Group (c. Beckeri Zone) of Kachchh (Spath. 1931:789), and they ma). perhaps also be found antong the supposed Torqrtatisphinctes and Poch?sphittctes of Collignon (1959) from hladagas,-. We have found only one or two specimens of Katroliceras (s. str.), K . porrinprri. below the Kimmeridgian-Tithonian boundary bed 8b. The principal Katrolic.~rosoccurrence in the Ken) a-Madagascar-Kachchh area (Ethiopian Subprovince) is therefore in the basal Tithonian Hybonotum Zone, and this occurrence may extend to Nepal (Enay, 1973). A significant member of this assemblage is Procrctspedircs cf. africutlus (Zwierzicky), which is available in two specimens. The one comes from locality Sf at the top of the section, together with H~botzoticerczscf. hyborlorurn;the other specimen was found by geologists of the British Petroleum-Shell Development Company at locality 1781, close to our locality 8e (Text-fig. 2 ) and. because of the horizontal attitude of the beds, at a very similar stratigraphic level. This second specimen was identified by the distinguished late W.J. Arkell (irl Eames and Clarke, 1964) with the earl) Cretaceous genus Groebericeras Leanza hitherto known only from Argentina. Consequently, the upper Changamwe Shales have been considered to be "Neocomian" in age (Walters and Linton. 1973). P. africanus is based on a single corroded specimen found loose by natives in the neighbourhood of the Smeei Bed of the Tendaguru Group in Tanzania (Zwierzicky, 1914: Janensch and Hennig, 1914). The similar Procraspeditcs ma:apilensis occurs in Mexico in the Glochirerasfialar Zone of the middle Kimmeridgian, subjacent to the H?bo~roriceras range zone, and thus approximately correlative to the Eudoxus Zone of Europe. Pachysphincres occurs throughout most of the section. It is based on P. africarlus Rertnanus (Dietrich, 1925) from the Kimmeridgian of Mahokondo near Tendaguru in Tanzania. We now believe P. germanus to be conspecific with P. be~richi(Futterer, 1894) from equivalent beds of the Mombasa area. The Phqlloceratina account for approximately 20 per cent and the Lytoceratina for a b u t 5 per cent of the total number of ammonoid specimens. taxonomic all^, they closely resemble the faunas of the H~bonoticerasrange zone of Madagascar and Kachchh. By far most abundant is Holcoph~lioceraspol~olcummesolcum (Dietrich) which occurs throughout the Hybonotum Zone and ranges downward at least into bed 8b. Dietrich's "species" from the (Lower) Kimmeridgian of Tanzania is here included in H. polwlcum (Benecke). According to Spath (1933:788, 791). the restricted species occurs above H. p. mesoleurn in the Katrol Group of Kachchh, but

this is not borne out by our assemblages, nor b~ Spath's listing in the same volume (1933:708). Only H. mesolcum [aff. polvolcun~(Benecke)] is listed from the "1st" and "2nd Katrol Assemblages", as well as from the Upper Tithonian ("Portlardian") Zamia Shales. Both .'species" were identified bj Collignon (1959) from : uppermost (Lower) Kimmeridgian "Zone of Aspidoceras longispinum and 70 quarisphincres alterneplicarus" of Madagascar. Of the less common "Mediterranean" Phllloceratinae. Ph~llocerus isor~puni Benecke has not been described from Kachchh or Madagascar: Culliph~l1oceru.i benacense (Catullo) is common in the "1st Katrol Assemblage" of Kachchh to which it was said to be restricted (Spath. 1933) and also occurs in the A . longispinurn and 7 alrerr~eplicarus Zone of Madagascar (Collignon, 1959): Pr~choph~llocer, p[vchoicum (Quenstedt). typically for the TethJan Tithonian, is also common in t: "1st" and "2nd Katrol Assemblages" of the H~bonorrcerasrange zone, and in thL "middle" and lower Upper Tithonian of Madagascar (incl. '.P. rithonicum", Collignon. 1959, 1960). auct.] is apparently Of the Lytoceratina, Lxtocerasfraasi Dacque [Hemil~tocera~ unknown outside of East Africa and of Madagascar where it occurs in the Upper Kimmeridgian (Collignon, 1959); while Ptero!vroceras monranum (Oppel) is we!! known from the Mediterranean Lower to Upper Tithonian and, probably, occur abundantly in the Kimmeridgian and lower Upper Tithonian of Manera 01, Madagascar (Collignon, 1960; cf. our synon>my). Lower Assemblage with Hybonoticeras pressuhm This not very abundant assemblage occurs directly subjacent to the upper assemblage described above. It includes Aspidoceras (s. srr.) spp. and the characteristic H?bonoriceras pressulum (Neumayr) in the middle and upper beds, and alsc Lirhacoceras? mombassanum (Dacque) in the lower and middle beds. The remainder of this assemblage is the same as in the upper assemblage. The small sample sizes may be responsible for the apparent absence of other taxa of the abundant upper assemblage, particularly since a number of them are known to range downward elsewhere (see above). This concerns particularly the basal 100 m of the section from which only Lirhacoceras? mombassanum has been clearly identified, while the specific identifications ofAspidoceras acanthicum (Oppel) and A. iphicerum (Oppel) are uncertain. These basal shales with horizon 8k can therefore be placed only in the Kimmeridgian. Approximately the upper 100 m (beds 8h-8b) of this lower sequence can be dated with some confidence b) the presence of H.vbonoriceras pressulum (Neumayr). This species has not previously been identified from this biogeographic province, although Collignon's (1959) "H. breisrofferi", supposedly from the Hybonotum Zone of Madagascar, is closely similar. Throughout its geographic range in Europe (Spain to Bulgaria), however, H . pressulum is confined to the lower Beckeri Zone, that is, the Subeumela Subzone of Sapunov (1977; see also Berckhemer and Holder, 1959: Enay and Geyssant, 1975; Banhel and Schairer, 1978). According to these authors, no H~bonoriceras is known from below the Beckeri Zone. Zeiss (1977, fig. 7) in Franconia, however, distinguishes the new zone of H . pressulum below the Beckeri Zone. We prefer to use the Beckeri Standard Zone in its broader sense and would include Zeiss's Pressulum Zone within the Beckeri Zone as a subzone.

Interval 8h-8b is therefore placed in the Beckeri Standard Zone based on the presence of H. pressulunl and its stratigraphic position subjacent to the Hybonotum standard Zone. Other apecies. for example. Pach?sphincres be~r-ichi(Futterer). to range almost throughout the local section (see under Upper Assemblage).

The uorldu ide differentiation of ammonoid fauna reached its peak in the late Jurassic a, rrjdenced b\ the provinciality of a varist! of organisms including belernnites. amn~onites.brachiopods, foraminifera. gastrop?ds. pelec\pods. certain coelenterates. and calpionellids. The ammonites have proved to be useful tools not only in ,tratigraphic correlation but also in delineating the patterns of provincialit!. Uhlig (1910) in his review of the works of Neumayr (1883, 1885). Nikitin (1886). and Haug (1907), distinguished four major realms. Subsequent authors recognized only three. the Boreal. the Tethqan, and the Pacific (Burckhardt, 1930: Arkell, 1956: lmlay. 1965) and, most recently, two major realms. the Boreal and the Tethyan. the latter including the Pacific Realm of other author3 (Stevens. 1963b. 1965, 1967. 1971: Hallam, 1969a. 1969b. 1971: Enay, 1973, 1980). The subdivision of the realms into provinces has varied similarly. Thus Arkell's (1956) Tethyan Realm was composed of the Mediterranean, the Himalayan, the Maorian. and the Ethiopian provinces, whereas Stevens (1967) considered the Ethiopian and Indo-Pacific provinces as derivatives of the Tethyan Realm (s. srr.): Enay (1973) formerly subdi\)ided the Tethyan Realm into the Mediterranean, the Himalayan (or lndo-Pacific), and the Ethiopian provinces, but recently (1980) only into the Mediterranean and Indo-Southwest Pacific provinces, the latter including the Ethiopian Subprovince. With the overwhelming acceptance of the plate-tectonic theory, recent authors (Stevens, 1967; Riccardi et al., 1971: Hallam, 1971: Enay, 1972; Verma and Westermann, 1973: Zeiss, 1974: Westermann, 1975; Westermann and Riccardi, 1976; Wiedmann, 1980) have projected their palaeontologic data on pre-drift reconstructions (particularly of Gondwanaland) of which several recent ones, based on geological and/or geophysical data, are presently available (e.g., Smith and Hallam, 1970; Dietz and Holden, 1970; Smith, 1971; Smith et al., 1973; Seyfert and Sirkin, 1973; Smith and Briden, 1977). Concerning Gondwanaland, these reconstructions differ mainly in the relative positions of Madagascar and the Indian subcontinent and in the timing of the plate movements. The most suitable reconstruction would be the one that offers the greatest compatibility among the geological, geophysical, and biogeographic data. Enay (1972) has plotted the Tithonian faunistic data on a global reconstruction that was modifieJ after Carey (1958) and resembles that of Smith (1971): this in turn is a combination of reconstructions by Bullard et al. (1965) and Smith and Hallam (1970). We agree with Enay that faunal realms should be viewed in the light of the actualistic palaeogeography taking into account the continental mobility. Enay (1972, 1980) distinguished during the Upper Jurassic the Boreal Realm ("Domaine Boreal"), the Tethyan Realm ("Domaine Tethlsien"), and, beginning only in the

late Tithonian, the Perigondwanian or Austral Realm ("Domaine Perigondwanien" The Boreal Realm consists of the Boreal Province which includes the lands presenr. bordering the Arctic Ocean (arctic Canada, northern Siberia. the Petchora Bas, Spitzbergen, etc.) and the Subboreal Province including the southernmost margins .)Streblites plicodiscus (Waagen, 1875:56, pl. X, figs. 5a, b) is probabl) an allied species but is poorly known (based on a single nucleus).

L%~l-~4j2

OM 34584 BOM 34585 IL11~-~~33

phragm ,.SMI.~p.34bod! ch.

W

HIW

76.0

c. 7.0

9.2

c. 39.0

c . 10.0

3.9

76.0 73.0 27.0

c . 5.0 c . 8.0 7.4

6.5 10.9 16.

41.0 39 0 16.5

c. 15.0

273

43.5

4.0

9.1

23.5

118.0

11.5

9.7

66.0

-

c.8.5 -

c . 12.0

1.94 -

5.5

superfamily Perisphinctaceae Steinmann, 1890 ~ ~perisphinctidae ~ i Steinmann, l ~ 1890 subfamily perisphinctinae Steinmann, 1890

Genus Pachysphincres Dietrich, 1925

Type Species Perisphincres (Pachysphincres) africogermatlus Dietrich, 1925 (by subsequent designation of Spath, 1931:467). Lectotype Here designated: original to Dietrich's pl. I , fig. 2 (copied in Arkell et al., 1957, fig. 427, la; Ib redrawn with altered final section). One of us (Westermann) has re-examined the type sets (autohyle) in the Museum fur h'atukunde der Humboldt-Universitat, East Berlin (courtesy of Dr. H. Jaeger who also supplied photographs), and came to the following conclusion. Among the several dozens of specimens in the type set from Mahokondo, Tanzania, are only a very few (including the smaller specimen figured by Dietrich) with "rounded broad whorls and deep umbilicus" to about 40 to 50 mm diameter, characteristic features of this species according to the original description. The large majority of the specimens (including the large specimen figured by Dietrich) have narrower inner whorls (depressed in nucleus only?) and a shallow-subconical umbilicus (and somewhat denser primaries, c. 25, vs. c. 20 per half-whorl). The best specimen according with the original description is designated the lectotype, that is, the medium-large original to Dietrich's pl. I , fig. 2 (copied by Arkell et al., 1957, p. L328, fig. 427, l a , b). Dietrich's whorl sections do not belong to the lectotype, but were probably taken from unidentified "P. africanus". The lectotype and all other specimens are, however, incomplete and strongly

,

distorted by "septarization" (diagenetic shrinkage in septarian concretions). lth shell displacement from the internal mould, subsequent calcitic infillings, etc.. ,li contributing to the difficulties in reconstructing the original shell. The type species is thus characterized by depressed, rounded, ovate, imr; . . whorls and very prominent biplicate ribbing with long primaries b e c ~ ~ : . , ; , ~ ataxiocerid on the outer whorl. The more complete. larger paralectotjpe (Dietrich. 1925, pl. 3. fig. I) and th, majority of the type set are perhaps identical with '.Perisphinctes afric.unus" Dacque, 1910, based on an incomplete septate specimen. Most specimens of the P africogermanus set, in fact. had been so labelled and Dietrich had realized :i, possible synonymy of his new name. He disregarded (?correctly), however, Dacc name as a notnen drrbirrtn, having been based on unsatisfactory material wit: ,, preservation of the important mature whorls. These specimens have also rather frequent simple ribs. Only accurate re-collecting at the type locality can establish the significance of the mentioned difference in the inner whorl and the affinities of "P. africanus". P. bejrichi (Futterer, 1898), described below, resembles the probable '.P africanus" discussed above. differing only in the somewhat more compressed ma1 whorls. We are using this name since it has seniority and was established on gi type material.

,

Comparison A closely related genus iskbrroliceras Spath, 1924, distinguished mainly in the bod) chamber with widely spaced. very prominent bullate primaries and four or five irregular secondaries per primary. The separation of these supposed genera, howeve,. remains dubious, since species with similar but regular Perisphincres-like multip secondaries also occur, for example, "Perisphinctes torquatus Waagen, (no11 Sowerby)" =Pachysphincres major Spath; as well as forms with finer secondaries and rather coarse primaries which Spath places in Karroliceras, for example, "K." arenosum Spath (and even inSubplanires, for example, S. adeloides Spath). Both of Waagen's species, "Perisphinctes" katrolense and "P. '' bathjplocus, are difficult to place and one has only to study Spath's (1931) monograph on Kachchh (e.g., pl 100, figs. la, b) to realize that even he admittedly could not clearly discriminate between many of the genera of his own creation. As usual he accompanieo genus-level splitting with species-level splitting, and the necessary taxonomic revision of this extensive fauna will be most difficult. The compressed, coarsely ornate forms of Katroliceras and Pachysphinctes are also hard to separate from Subdichotomoceras, particularly in the microconchs of the former two where the coarse biplicate ribbing typical for the latter may continue almost to the aperture (many examples in Spath, 1931; see under K.? aff. bathxplocus). We have considered relatively shon primaries and coarse ornament of the inner whorls as distinguishing characteristics forSubdichotomoceras in this case. It should be remembered, however, that the type species of this genus is from Boreal central to northern England and the Tethyan form may be a homeomorph allied to the Pachjsphincres-Katroliceras group. This comes to mind when studying the co-occurring P . beyrichi and S . ? cf. sparsiplicarum from Mombasa. This problem can only be solved with excellent material from both faunal realms at hand. Originally, we placed Pachjsphinctes as a subgenus in Katroliceras, but we have

in separated them genericall) because of the very marked differences of the type at maturity. Again, a thorough re-examination of the entire species complex is @ded. ~~~~h~~ apparently related, poor11 known and often nlisinterpreted taxon is Torqyl~spl~i~~ires Spath, 1924. The type species, Irnoa~~iires rorqirarus J de C . lowrby (1840). is based on a rather poor11 preserved, small. and incomplete holotype ("figured by Spath. 1931. pl. 76. figs. 4a.b) which ma) be either an inner of a much larger macroconch or an almost complete microconch. The typically ,,lmu1ate shell has subquadrate whorls bearing dense ribbing which is distinctive in (br preynce of abundant simple ribs. Dr. Jai Krishna (pe" comm.) has collected what he considers to be topotypes from the Kimmeridgian of Jaiselmer in Rajasthan. incolnplete small specimens of modest preservation all shoa the densely ribbed subquadratic whorls with abundant simple ribs. differing markedl) from k'orrolireras and other similar coarsely ribbed forms. Torqrwtisphinctes. however, remains poorly known so that the generic name should be used only with extreme caution. Another genus, well known from its type locality at Niti in the Himalayas, and p b a b l y more widely distributed than is apparent from the descriptions. is ~ulacospliir~ctoides Spath, 1924. The type species, Aulacosphinctes ir~fundihulus Uhlig (1910). as well as a host of similar and probably intergrading "species", are characterized by depressed elliptical to ovate whorls with mostly dichotomous dense sigmoid ribbing, often followed by a triplicate and/or ataxiocerid adult stage with stiffer primaries. Thus, it is distinguished from Pachysphinctes by the more inflated whorls and, typically, by the more sigmoid ribbing, but the distinction becomes spurious with intermediate forms and both are certainly closely related. We retain these taxa at the generic level only because we are unable to solve the taxonomic problems at the genus-group level without re-examining all the type material and sections. We follow Dr. John Callomon (pers. comm.) in transferring the entire group from the Virgatosphinctinae to the Perisphinctinae because of the close similarity of some of these forms, for example, Karroliceras and Pachysphinctes, to typical Perisphinctes with the exception of the adult body chamber. Iga

,

Pachysphinctes beyrichi (Futterer, 1894) PI. 6, figs. 1, 2a.b. Text-fig. 4 Perisphincres bevrichi Futterer, 1894:9, pl. 2, figs. 1-3. Perisphinctes (Virgarosphinctes?) beyrichi-Dacquk, 1910: 14, pl. 4. 2a.b. ?Perisphinctes africanus Dacqut, 1910: 17, pl. 3, fig. 2. ?Perisphincres latissimus Zwierzicky, 1914:65, pl. VlII, fig. 4. ?Perisphinctes (Pachxsphinctes) africogermanus Dietrich. 1925: 12, text-figs. ? I , ?2, and pl. 3, fig. 1 only. T0rquarisphinctes bevrichi-Spath, 1930a:55, pl. 3. fig. 6 [cum synonymy; lectotype des. Futterer, 1894, fig. 21. ?Perisphincres cf. denseplicatus-Dietrich. 1933:20, pl. 1, fig. 1. '?TorquatisPhinctes betsibokensis Collignon, 1959, pl. CIX, fig. 401. ?Subplanires adeloides Spath---Coliignon, 1959. pl. 127, fig. 475.

Material Four. possibly seven specimens.

ROM

34624, damaged. almost complete; from lot.

8b. K N M I - ~ ~small 3 5 , specimen with one-third whorl body chamber well pre,:.-. j , ?ROM34620. ?ROM34746. ~ ~ ~ 1 . 5 1 4large 3 6 . specimens with partial body ch, from Ioc. 8d. ?ROM34605. whorl fragment: from loc. 8h. ROM 34733, crush^, . 1 incomplete: from loc. 8j. Description The shell is typically planulate u ith wide and shallow but srepped umbilicus. The whorl section changes from moderately depressed ovate at the nucleus at 20 to 35 r.;, diameter (U'IH" 0.65), to rounded subtrapezoidal or subquadratic in their larger whorls. The ornament consists mostly of moderately dense, high, and sharp b i p l ~ ~ a r c ribbing with a few simple ribs: additional secondaries, however. appear in the adult stage on the outer whorl greater than 100 mm diameter, beginning low on the flank and producing ataxiocerid ribbing. The primaries are prorsiradiate on the nucleus bur rectiradiate on the outer whorls, while the secondaries cross straight over the venter There are also about three oblique constrictions per whorl. The septal suture is rather simple, with deep E, slightly shorter L, and se\ much smaller, oblique, and retracted umbilical elements. Remarks The lectotype of "Perisphirlcres" beyrichi Futterer (here refigured, Text-fig. 4) from the ?Kimmeridgian of the Changamwe Shale near Mombasa is a large fragmentac macroconch with an inferred diameter of about 170 mm, while the paralectotype !\ septate to the end of 71 mm. The intermediate whorls of both specimens (60 70 mm D) are rounded, subquadratic, slightly depressed, with steep umbilical wa. . and bear 22 to 33 prominent, somewhat prorsiradiate, and partly flexed primaries per half-whorl. While biplication is the rule, rare single ribs and very scarce trifurcation also occur. Trifurcation with irregular fasciculation (ataxiocerid ribbing) is clearl! present on the outer whorl of the lectotype, while the secondaries of its penultimate whorl are not visible. The septal suture also agrees with Dietrich's illustration (1925. text-fig. 1) of supposed P. africogermanus and our specimens. Although Dietrich (1925) did not even compare his P. africogermanus with P bewichi, we suggest that the abundant planulate form (not the lectotype) that he placed in his species is probably conspecific with P. be~richi. Specimens closely resembling the lectotype of P. be~richioccur rarely in the Beckeri Zone, and also in the Hybonotum Zone where they appear to intergrade into more coarsely ribbed forms, that is. 19 to 21 versus 22 to 24 primaries per half-whorl. The coarsely ribbed forms (loc. 8d) with rather shon primaries and occasional simple ribs, may perhaps belong to a distinct subspecies. However, no complete specimen has been found and some of the largest specimens may not be fully grown (last septa not approximated). Collignon (1959) illustrated two similar but poorly preserved "species", respectively from the Hybonotum Zone and Kimmeridgian of Madagascar, under "Subplanires adeloides Spath" and "Torquatisphincres bersibokensis" Collignon. Stratigraphic Occurrence Locally abundant in the Hybonotum Zone (mainly coarsely ribbed variety; ?subsp.): scarcer in the Beckeri Zone (s. srr. ), and probably also lower.

ucre lo\[. ( D r . J . Helmz. per,. comm 1

hleasurernents SPEC.I\IE\\O I;\vI-M~~~ hod! ch

end p h r a f m .

K\\llL\l435

hid!

ch. rhrasili plira;ni phrdfn' phrssm

C

UID5

H

M'

HIW

Genus h'afroliceras Spath, 1924

Type Species Arnrnonires porringeri J. de C. Sowerby. 1840 h'afroliceras poningeri (Sowerby, 1810) PI. 4, figs. 1a.b An~moniresporrirtgeri Sowerhy, 1840:719. pl. 61, fig. 10. (7) Perisphincres porringeri-Waagen, 1875: 183. pl. LI, figs. la,b [ = Karrolrc ,, ~t.aageniSpath, 193 1 :508]. Perisphirlctes pofringeri-Futterer, 1894:7, pl. 1, figs. 1, 2. Karroliceraspottingeri-Spath. 1930a:56: Spath, 1931 :505, pl. CII, fig. 5 [Futterer. 18941: pl. LXXXIII, fig. 4; Collignon, 1959, pl. CXXI, figs. 455, 456. (?Karroliceras ulaageni Spath, 1931 :508 [for Waagen, 18751. (1)Karroliceras lerertse Spath. 1931:511, pl. LXXXIX, fig. 1. Karroliceras porringeri Souerby var. onala~~e/onensis Collignon, 1959, pl. CX . fig. 457.

.

,LA,.,>>

-

.-

.,

c~unc.! ,>f Dr. M . K. Houanh. Br~tisnMu\eum. 3alural

H15ton

New Material Two or three specimens, more or less strongly crushed andlor distorted. ROM 34749, 37, with complete specimen with body chamber: from loc. 8c. K N M I - ~ ~phragmocone bod) chamber; from loc. 8h. ?ROM34748, fragment of body chamber, from loc. R Remarks The additional number of "species" erected by Spath (1931) on the Kachchh fauna is obviously excessive, and we have here made a tentative attempt at placing some of these into synonymy. For proper taxonomic revision, however. new collections from the exact stratigraphic levels are necessary, as is the examination of all existing material. As a result of Spath's taxonomic splitting, no good specimen of the "true" K. katrolense (smsu Spath) from Kachchh has been figured photographically. The only complete, but exceptionally small specimen figured by Spath (1931, pl. 102) and in the Treatise (Arkell et al., 1957, fig. 425) under that name is Futterer's old specimen from Mombasa. Futterer ( 1894) compared his Kenyan "Perisphincres porringeri" with Waagen's specimen from Kachchh which Spath (1931) renamed Karroliceras ufaogeni. Futterer's specimen, however, is a trueK. porringeri. Spath (1930a) did not figure his "true" K. porringeri from the Mackinnon Wood collections. Our material has yielded examples which are unquestionably identical to Sowerby's and probably Spath's poorly illustrated Kachchh forms. "K. waageni" Spath is said to be distinguished by the depressed subrectangular inner whorls bearing long primaries, but is here tentatively included in K. poffitlgeri, together with several others of Spath's supposed new species. The holotype of K. pofringeri from Kachchh has apparently never been refigured. It is illustrated here (Text-fig. 5), courtesy of Dr. M.K. Houranh, British Museum, Natural History.

A good series was illustrated b) Collignon (1959. fig\. 155-57. 7 4 5 8 4 1 ) from the HYbnotum Zone of Madagaaar. Collignon. however. a p p e r s to hare erected an unduly large number of species on h s materid: nian! are based onl) on inner u'horls which are similar in many species. The wedge-hhaprd primaries that usually characterize Karroliceras do not make their appearance until a fairl) late stage in pwd. so that coniparisons hetween species of this genus should l x b a a d upon adult sprcimens or those in which the characteristic body-chambcr ornamentation has started to appear

Stratigraphic Occurrence Rare in Beckeri and Hybonotum Zones. Measurements SPECIME\ NO KNHI.\1h?7 ROM 34748

D 137.0 133.0

IJ

77.0 710

U/D% I

.

0.53

H

W

HiW

36.0 39.0

43.0 49.0

0.79

o.88

Material 8. complete uith pmlal h d ! One hpecimen. K U M I - ~ ~ 3nearl! loc. 8c.

chamber. from 43

Description The shell is large. stoutly planulate. with depressed subrectangular whorl The umbilicus is wide and shallow with almost vertical umbilical wall. The LLidr,e primary ribs arise at the umbilical shoulder, are rursiradiate at the umbilical curve forward on the umbilical shoulder, continue prosiradiate over the subparallel flanks, and thicken at the ventro-lateral margin. Up to a diameter of 110 mm the primaries give rise to convex secondaries traversing the venter. At large diameter\, the primaries trifurcate with an occasional intercalatory Irregularly placed fcrbie constrictions are present. Remarks This form is characterized by its large size and the broad, stout whorls. The long coarse primaries probably did not quite reach the wedge-shaped distant stage as I" typical K . portit~geri,but the body chamber is incomplete. This specimen agrees with Waagen's form, named K . w.aagetli by Spath but considered by us to be prohabl! conspecific with K. porringeri. K. katrolerzse (Waagen) has similar ribbing 11 much less stout. The Madagascan "Karroliceras pingue Spath" describe, Collignon from the Hybonotum Zone (1959, fig. 461) is similar but incomplete.

p m i n e n t , somewhat prorsiradiate to forward-curved primaries. The outer whorl biplicate ribs, except for rare s~mpleribs. and two additional secondaries are on the body chamber. The aperture has a deep constriction, ventral flare, and p" right lateral lappet preser\'ed at 12 1 mm diameter. ourspecimen differs from K . ? bath~p\;plocusin the weak adult modification of the The primaries continue essentiall) unmodified but slightl) more prominent @,he apnure. while only two triple divisions are developed. The figured syntype of waagen hasan incomplete body chamber of at least 130 mm diameter but already &wS strong modification of ornament toward the end with expanded spacing of e k e s and several intenaiatory and fasciculate secondaries Therefore, we lentatit,el) transfer this species to K a f r o 1 i r . c ~ ~Spath's ~. specimens of "P." ~ r h ~ p i o r u and s "P." linguiferus also shou abundant extra secondaries and expanded spacing of the primaries on the ultimate half-whorl and both appear to be identical with Waagen's species. Our specimen is therefore somewhat intermediate between Katroliceras, Parhysphinc.les. and Subdichotomoceras indicating taxonomic splitting, but we belive it to be a paeJomorphic variant or subspecies of K.? barh~plocus.A number of similar forms, including this species, also occur in the H)bonotum Zone of Madagascar (Collignon. 1959).

*

Stratigraphic Occurrence Stratigraphic Occurrence Beckeri Zone.

Rare in Hybonotum Zone.

Measurements SPEC~MEN NO

D

U

U/D%

H

W

HIW

K~MI-~~38

131.0

65.0

49.6

40.0

56.0

0.71

Measurements SRCIME~ \a

D

U

U[D%

H

W

HIW

121.0 85.0

57.6 39.5

47.0 46.0

c. 30.0

c. 36.0

21.0

27.5

c . 0.85 9.75

KNMI-MA~~

apenure

Katroliceras? cf. K. bathyplocus (Waagen, 1875) $ PI. 5. figs. 2a.b ?Perisphincres bathjplocus Waagen, 1875: 192, pi. 50, figs. la,b. ?Pachysphinctes bathylocus-Spath, 1931:493, pl. 77, fig. 1; ?pl. 93, figs. 5, 9: p 96, fig. 4. ? ~ a + ~ h i n c ? e s Iinguiferus-Spath, 1931:496, pl. 97, fig. I; p l 98, fig. 5. ?Pachysphinctes linguiferus-Collignon, 1959, pl. 118, fig. 446. ?Pachysphinctes barhyplocus Waag. var. sparsicosrata-Collignon, 1959, pl. 118. fig. 447.

Material One, possibly two microconchs, partly with base of lappets, KNMI-~A39,slightl! damaged with somewhat crushed body chamber; from loc. 8h. ?ROM 34615, more strongly crushed and inner whorls obliterated; from loc. 8b. Remarks Our figured specimen closel) resembles Waagen's species from the "Middle Katrol Beds" of Kachchh, in particular also the excellent microconchs figured by Spath (1931) under K . bathyplocus and K. linguiferus. The inner whorls bear rather dense

end phrapm

Genus Subdichotomoceras Spath, 1925

Type Species

S. lamplughi Spath, 1925. Remarks The t!pe species is from the Lower Kimmeridgian (s. angl.) of Yorkshire. England, AutissiodorenSis Zone (Cope, 1978), correlative with the Beckeri Zone. This is an evolute planulate form with subquadrate whorls, characterized by the ver). prominent biplicate distant ribbing with rather short primaries, persisting to the adult body chamber and also on the inner whorls. The genus has usually been considered cosmopolitan in distribution (e.g., Arkell. 1956. 1957), but doubts have more recently been expressed about the affinity of the Tethyan forms which ma) be homeomorphs (e.g., Enay , 1973). A brief examination of L.hlig3s (1903, 1910) work on the Spiti Shales, Himalayas, and of Spath's (1927-33) work on the Katrol Beds of Kachchh, western India, suggests that some

Description The shell is large, stoutly planulate, with depressed subrectangular whorl The umbilicus is wide and shallow with almost vertical umbilical wall. The primary ribs arise at the umbilical shoulder. are rursiradiate at the umbilicd hall, curve forward on the umbilical shoulder, continue prosiradiate over the subparallel flanks, and thicken at the ventro-lateral margin. Up to a diameter of 110 mm the primaries give rise to convex secondaries traversing the venter. At large diamears, the primaries trifurcate with an occasional intercalatoly, Irregularly placed feeble constrictions are present. L

~

Remarks This form is characterized by its large size and the broad, stout whorls. The long. coarse primaries probably did not quite reach the wedge-shaped distant stage as in typical K. pottirigeri, but the body chamber is incomplete. This specimen agrees with Waagen's form, named K. waagetli by Spath but considered by us to be probabl! conspecific with K. potfingeri. K. katrolerlse (Waagen) has similar ribbing b much less stout. The Madagascan "Katroliceras pingue Spath" describe< Collignon from the Hybonotum Zone (1959, fig. 461) is similar but incomplete.

,

prnminent, somewhat prorsiradiate to fomard-curved primaries. The outer whorl biplicate r i b , except for rare simple ribs. and two additional secondaries are ,,I on [he body chamber. The aperture has a deep constriction, ventral flare, and P . lateral lappet preserved at 121 mm diameter. right differs from K . ? bath?plocus in the weak adult modification of the our ,,,,,merit, The primaries continue essentiall) unmodified but slightl) more prominent m,bapenure, while only two triple division, are developed. The figured smtype of Waagen has an incomplete body chamber of at least 130 mm diameter but alread) Strons modification of ornament toward the end uith expanded spacing of ,@ and several intercalatory and fasciculate secondaries. Therefore, we untativel) transfer this species to Karroliccrus. Spath's specimens of "P." ~ , h ~ p l o c uand s "P." linguiferus also show abundant extra secondaries and spacing of the primaries on the ultimate half-whorl and both appear to be idintical with Waagen's species. Our specimen is therefore somewhat intermediate between Kurroliceras, PachjSphincres, and Subdichotomoceras indicating taxonomic splitting, but we belive it to b: a paedomorphic variant or subspecies of K . ? bath~plocus.A number of similar forms, including this species, also occur in the Hqbonotum Zone of Madagascar (Collignon. 1959)

~

~

~

*

, ,

Stratigraphic Occurrence Rare in Hybonotum Zone.

Stratigraphic Occurrence Beckeri Zone.

Measurements Measurements SPECIMEN YO KNMI-MA39 apenure Kafroliceras? cf. K. bathyplocus (Waagen, 1875)8 PI. 5, figs. 2a,b ?Perisphinctes bath~plocusWaagen, 1 875: 192, pl. 50, figs. la,b. ?Pach\.sphinctes bafhyplocus-Spath, 1931:493, pl. 77, fig. I; ?pi. 93, figs. 5, 9; p 96, fig. 4. ?Pachjsphinctes linguiferus-Spath, 1931 :496, pl. 97, fig. 1; pl. 98, fig. 5. ?Pachysphinctes linguiferus-Collignon, 1959, pl. 118, fig. 446. ?Pachwphinctes bathyplocus Waag. var. sparsicostata-Collignon, 1959, pl. 118. fig. 447.

Material One, possibly two microconchs, partly with base of lappets, KNMl-M~39,slightlj damaged with somewhat crushed body chamber; from loc. 8h. ?ROM 34615, more strongly crushed and inner whorls obliterated; from loc. 8b. Remarks Our figured specimen closely resembles Waagen's species from the "Middle Katrol Beds" of Kachchh, in particular also the excellent microconchs figured by Spath (1931) under K. bath~plocusand K. linguiferus. The inner whorls bear rather dense

end phragm

D

U

UIDQ

H

W

HIW

121.0 85.0

57.6 39.5

47 0 46.0

c 30.0

c. 36.0

c . 0.85

21.0

27.5

9.75

Genus Subdichotomoceras Spath, 1925

Type Species S. lamplughi Spath, 1925. Remarks The t!pe species is from the Lower Kimmeridgian (s. angl.) of Yorkshire. England, Autissiodorensis Zone (Cope, 1978). correlative with the Beckeri Zone. This is an evolute planulate form with subquadrate whorls, characterized by the ver). prominent biplicate distant ribbing with rather short primaries, persisting to the adult body chamber and also on the inner whorls. The genus has usually been considered cosmopolitan in distribution (e.g., Arkell, 1956, 1957). but doubts have more recently been expressed about the affinity of the Tethgan forms which ma) be homeomorphs (e.g., Enay, 1973). A brief examination of Lhlig's (1903, 1910) work on the Spiti Shales, Himalayas, and of Spath's (1917-33) work on the Katrol Beds of Kachchh, western India, suggests that some

Karroliceras have septate whorls similar to the more depressed f,>. Suhdichoromoceras and that some coarsel! ribbed Aulacosphinctoides are \ L . to relati~el)finel) ribbed Subdichoto~noceras. Difficulties also arise in the scnL,,, separation from Karrolicerms of microconchs with persistent biplicate ribbing. u,,h perhaps one or two simple ribs or additional secondaries. Is the absence of the adu]! modification, so characteribtic for Karroliceras (and Pachysplrincres?), simp11 due mlcroconchiate, progenetic shortening of the ontogeny, or is the persistent biplicarlon diagnostic for Subdichotomoceras? What taxonomic weight is to be attached ,.' inner whorls versus the bod) whorls? Obviously, Spath (1931) had th. d~fficultieswhen confronted with the Katrol fauna as is evident from examln ., monograph (e.g.. pl. 82, figs. 1 , 2).

r.an -- tb H!bonotum

Zone of Madagawnr. a poorl! hnoun form based on a uhorl fngmsnt ""I!. ours F ~ ~ m e n(and s Z u ~ e r z i c k )h ' ~n l )appear to d ~ f f e from r the slightl) younger ,,%hchh 'pecies (Spath. 1931) b) the more rounded uhorl section uithout a lateral sbulder. although this could be caused b! crushing in the other specimen,. stratigraphic Occurrence

H) Mnorum Zone.

;

$teasurements SRClME'.

Subdichotomoceras (?) cf. S . sparsiplicatum (Waagen, 1875) PI. 6, fig. 3; PI. 7, figs. l a , b; 2

50

end phragm-i h e . hod) ch. phraorn

D

U

UIDQ

H

W

HIW

c . 129.0 c 84.0

66.0 c.410

c . 51.0

c 38.0

c . 39.0

c.490

c.26.0

c32.0

c. 0.97 c.0.81

?Prrisphi~rctessparsiplicur~rsWaagen, 1875:204, pl. XLIX, figs. 2a,b. Perisphinctes sparsiplicarus-Zwierzicky , 1914:61, pl. VIII, figs. 1,2. ?S~thdichoro~noceras sparsiplicatrtm-Spath, 1931 5 2 3 , pl. 82. fig. 7; pl. 86, fig. 7 Subdichotomoceras? aff. S . sparsiplicatum (Waagen, 1875) PI. 5, figs. la-c

Three specimens. K N ~ I I - M A entirely ~ ~ ~ ~ . septate and panly crushed; KNMI-M440: from loc. 8d. KNMI-M.47731,septate or with less than one-third whorl body c h a m k r part11 crushed; from loc. 8c. Material Description The large shell had evolute and probably somewhat depressed (now partially crushed) ovate to subelliptical whorls. The ornament of the phragmocone consists of biplicate. extremely prominent ribs, with 13 to 16 primaries per half-whorl, at least as far back as 20 mm diameter. The primaries are recti- to somewhat prorsiradiate and divide 31 or sllghtl) above mid-flank into pairs of radial secondaries. These are only sligh less prominent than the primaries and pass straight over the venter. The phragmoco. . of two specimens was at least 120 mm in diameter. One specimen (PI. 6, fig. 3) has minute lateral tubercles on the intermediate whorls. The ultimate phragmocone whorl was about as high as broad. The body chamber is unknown except for its beginning which resembles the phragmocone. One or two deep, oblique, and continuous constrictions per whorl are present on all specimens, usually followed by a simple rib. The septa1 suture is known only wit': umbilical elements which are moderately complicated, strongly oblique, an retracted. Discussion According to Spath (1931 ), S.? sparsiplicarum occurs in the Middle Tithonian uppermost Katrol Beds of Kachchh, not in the basal ones as originall) assumed b) Waagen (1875:205). The species also occurs in the Lower Tithonian Smeei? and Nerinella beds of the Tendaguru Series in Tanzania, together with Pnch~sphincres be~richi(Futt.) (Zwierzick). 1914; Dietrich, 1925, 1933; Arkell, 1956). This form resembles Collignon's (1959, pl. 127, fig. 478) "Dorso~lanites?ar~rsalovensis"

One specimen. K N M I - ~ ~ almost 4 1 , complete without aperture; from loc. 8b Description The shell is planulate, slightly involute with a wide and shallow umbilicus. The umbilical wall is rounded and steep. The whorl section is semicircular with weakly curved flanks and broadly rounded venter. The ornament consists of thin, rather dense primaries which arise at or near the umbilical shoulder. Most of the long primaries bifurcate into very prominent secondaries while a few remain single. Both primaries and secondaries are projected slightly prorsiradiate crossing the venter somewhat convexly. On the body chamber a single trifurcation is also present. Constrictions are present throughout growth but are more distant in the early whorls than in the latter on account of the normall) deep intercostal spaces between the primaries. Remarks While the phragmocone resembles that of the relatively compressed Karroliceras ex. er. karrolense (Waagen) as well as of Pach?sphincres, the body chamber has the biplicate. v e v broad, and prominent ribbing of Subdichotomoceras. This form, therefore, differs from P. sparsiplicatum in the longer primaries and the denser ribbing of the inner whorls. Stratigraphic Occurrence Rare at boundaq of the Beckeri and Hybonotum Zones.

Measurements

Lithacoceras? mombassanum (Dacque, 1910) PI. 8. figs. 1a.b

Family Ataxioceratidae Buckman, 1921

Genus Lithacoceras Hyatt, 1900 Type Species Amnloriires ul~nettsisOppel, 1858 (cf. Berchhemer and Holder, 1959, fig. 501 Remarks The Lithacoceratinae Zeiss, 1968 are included in the Ataxioceratinae. Discosphiric.r~~~ Dacque and Subplanires Zeiss respectively were considered to incorporate the L-ppi.1 Oxfordian and Kimmeridgian-Lower Tithonian microconchs of Lithacoceras b) : (1966) and Zeiss (1968). The type species of the former, D. arussiorum Dacque Somalia, is, however, poorly known and its stratigraphic occurrence needs to b~ reinvestigated. The type species of Subplanites. S. reisi (Schneid), appears to be distinguished from D. arussiorum essentially in the ataxiocerid (simple virgatotome~ ribbing of the adult body chamber or its terminal portion only. We do not consider the existing evidence strong enough to draw the apparent dimorphs into synonymy, bur tentatively include the apparent microconchs in the genus Lirhacoceras (s. I.).

.Poisphi,lc.re5 -

(lirgarosphincres) mornbassanus Dacque. 19 10: 15. pl. 111. fis. 4: pl. IV. fig. 1 -,ifhocflceras rt~ombassar~um-Spath. 1930a: 48. pl. I\.'. flg. 1 t lector! pe des.: Dacqu& 1910. fig. 1). ofhacflccras rr~ackirtnor~-~r~oodi Spath, 1930a:49, text-fig. 2. ~i,hacocerosroub~anurn-Spath, 1930a:5 1. tezt-fig. 3. ?rjfhacoceras rorquariforme Spath. 1930a:49. pl. 4. fig. 14 Material Three to fi\.e specimens. KNh11-~~43a, slightl) crushed. large and almost complete; from Ioc. 8k: ?ROM34734 and ?ROM34736, incomplete phragmocones: from locs. 8a and 8c; ROM 34737, external mould of partl! crushed specimen; from loc. 8h. ~ N M I - Mwell-preserved A ~ ~ ~ , juvenile(?); from loc. 8d. Remarks This species is said to be one of the most common and highlq variable form5 in the Kimmeridgian of Changamwe (Spath. 1930a). We nevertheless include in this species also L. rnackinnon-naoodi Spath, his ..L. roub~arlunz" (non Fontannes), and perhaps also his L. rorquariforrne, since Spath (1930a:48-50) admitted the presence of transitional forms. Stratigraphic Occurrence

Lithacoceras cf. L. fraasi (Dacquk, 1910) PI. 8, fig. 2

The species is common in the Kimmeridgian of Mombasa, and may extend into the Hybonotum Zone.

?Perisphirlcresfraasi Dacquk, 1910: 20, pl. 4, fig. 3, text-fig. 10 (lectotype). ?Lirhacflcerasfraasi-Spath, 1930a:52, pl. 3, fig. 1 ; pl. 6, fig. 2; pl. 7, fig. 5. ?Lirhacoceras fraasi--Collignon, 1959, pl. 103, fig. 386. Material One Specimen. KNMI-MA42, partly crushed and incomplete; from loc. 8c Remarks The species was based only on a small septate specimen (the lectotype) and a fragment. Spath's (1930a) specimens from the Kimmeridgian of Changamwe were also all fragmentary so that the species remains poorly known. Our specimen remain. densely ribbed to the preserved end at 114 mm diameter. Collignon's (1959 specimen from the early Kimmeridgian (pre-Beckeri Zone) of Madagascar is also com~lete. This species is characterized by the small umbilicus and the persistence of dense ribbing. Stratigraphic Occurrence The species seems to range through most of the Kimmeridgian into the basal Tithonian. Our specimen is from the Hybonotum Zone.

Lithacoceras cf. L. albulum (Quenstedt, 1887) PI. 8, fig. 3 ?Ammonites planularis albulus Quenstedt, 1887: 1076, pl. 125, fig. 8. ?Perisphincres (Lirhacoceras) albulus-Berckhemer and Holder, 1959:55. pl. 9, fig. 48; pl. 10, fig. 5 1 (holotype refigured); text-fig. 32b. Material One specimen. K N ~ ~ I - M large A ~ phragmocone ~, and half-whorl damaged body chamber u ithout aperture; from loc. 8b. Remarks According to Berckhemer and Holder (1959:55) and Zeiss (pers. comm., 1978). L. albulum occurs onl) in the Hybonotum Zone of Swabia and is characterized by the early appearance of virgatotome multiple secondaries and relatively prominent primaries, and by inclined constrictions. Our specimen closely resembles the Paratype figured by them as figure 51, while the holotype has finer secondaries.

Stratigraphic Occurrence

JO

Rare at boundary of the Beckeri and Hjbonotum Zones

!

60

Genus Phanerostephanus Spath, 1950

Type Species P. s u b s e ~ c rSpath, 1950.

.-

phanerostephanUS (Nothostephanus) d i d p. (N.) kurdistanensis p . (p.) subsenex P. (P.) hudsoni p. (p.) intermedius

Discussion on the bah15 Spath (1950) created the genera Phanerosrephanus and h'orh~sre~hanus of Upper Jurassic material from Jebel Gara near Amadia in Kurdistan, northern Iraq He placed Phanerosrephanus in the subfamily Virgatosphinctinae and hlothostephanus tentatively in the Virgatitinae, but Arkell et al. (1957:L330) considered both "genera" as Virgatosphinctinae. Phanerosrephanus included P . subsenex ( species), P. hudsoni, P . intertnedius, and P. dalmasiformis, all described by Sy Nothosrephunus, however, was based on the single, and so far only known, spec,?\ N. kurdisranensis Spath, with unknown body chamber. The closer examination of the "genera" reveals several similarities. Both are compressed platycones with depressed circular whorl sections in the early whorls, giving rise to high whorl sections with arched venters in the later whorls. Virgatotome to fasciculate ribbing characterizes the early whorls including the main portion of the phragmocone, g i \ ~ n c rise to broad umbilical tubercles and smooth flanks in later whorls including the bc chamber. Both have irregularly spaced, shallow constrictions. The septa1 suture nearly identical, the adult suture consisting of a broad external saddle, a trifid lateral lobe, which is almost as deep as the external lobe or deeper, and two more saddles on the whorl side (Spath, 1950: 104, 115). The only marked differences between the two "genera" are the degree of involution and the growth stage at which the ornament transition occurs. Both features, however, vary intraspecifically. By and large. the adult shell of Norhosrephanus kurdisranensis is involute; th umbilical width of the holotype and of other specimens figured by Spath (1950, pl. 7 figs. 1-4, 8) ranges from 20 to 25 per cent of the diameter. The umbilical widths oi Phanerostephanus subsenex, P. hudsoni, and P . intermedius range from 35 to 47 per cent (Text-fig. 6). Measurements of the better-preserved specimens of P. ( N . ) digoi sp. nov. indicate that it has an enlarged umbilicus in the outer whorls even after the uncoiling normally observed in the last whorl of adult ammonites is taken into consideration. The umbilical width of the inner phragmocone whorls ranges from 19 to 26 per cent. In adult specimens, however, uncoiling proceeds (29 to 41%. Text-figs. 6, 7). This appears to indicate that the new form described here includes the range covered by both Norhosrephanus and Phanerostephanus. With regard to the ornamentation, it should be pointed out that No~hostephunusis known mainly from the phragmocone. The bolotype has only a very small portion of the body chamber which is crushed (Spath, 1950. pl. 7, fig. la). Presumably on the basis of the other material in his collection of about 30 fragmentary and crushed specimens, Spath (1950: 115) stated that "the secondary ribs seem to disappear on the body chamber and probably only the blunt and rounded umbilical tubercles remain." 7

-

,.,.,. ,,,,, ,,.,

T ~ , ~ . ( ,6~ vmhillcal

,,,

..,...

l ~ u ~ l .P~, ~( p~ ,1, I ~I I~~ ,
with the ornament of G. irius. The European stratigraphic distribution of Gra\,esio is restricted to thL T~thonian Hjbonotum Zone in which, in contrast to earlier opinions. :. : :r .e recogmized species appear to occur either in association or in time-equi\aisnt &J, (Banhel, 1959; Hahn, 1963: Cope and Zriss. 1961: Enay. 1966. 1973: Zeiss. 1 9 6 ~ ~ The occurrence of Grot,esia also in East Africa (and ?Madagascar) is ren~arl,~hl, bscause the other occurrences are restricted to western Europe, that is. to France, German). and southern England (op. cit.), and perhaps also to the Rubsian plaiti,rm ,

and western Siberia (Zeiss, pers. comm.). Perhaps the disjunct geo.. . distribution is due to the unusually strong ecological restriction of Grat,e~ic. : &iss (1968: 152) considers to be the inner sublittoral (c. 10 to 40 m depth): th, >~.i.,, occurs in Europe in similar association with perisphinctids, aspidoceratidb. arid H~bor~oriceras. The credit for the significant discovery of Gra\,esiu in East Africa should ,UO to Arkell who identified specimen RGM 1891 (British Petroleum-Shell De\'elopnlrn: Company's collections) with Gruvesia cf. G. portlut~dica(de Loriol). That spei.i:\ ,, no& regarded as ajunior synonym ofG. gigus (Hahn, 1963:98: neotype pl. 9. t Arkell's identification in Linton and Maclean's report (1955). however, ren.. unpublished. To our knowledge the only previous record of Gravesia outside Europe and western Siberia is that of Collignon who figured a minute specimen (D =21 mnll from Madagascar as Gra~zesia?sp. indet. (1960, fig. 647). Because of its doubtful identity. Hahn (1963) and Enay (1966) correctly regarded this occurrence as poor evidence for the distribution of Gravesia.

..

Stratigraphic Occurrence Rather common in the Hybonotum Zone Measurements SPECIMES NO

KNMI-~~48

Holotjpe end bod! ch. end phragrn. KNMI-MA49

end phragrn

ROM

348 19

Gravesia aff. G. loupekinei Sp. nov. Q PI. 13, figs. l a d

~pterial Ow spclmen. KNMI-MASO. well-preserved phragmocone, siightl) damaged internal puld with test remains: from loc. 8e. Description m i s large specimen is still septate at 132 rnm diameter. The shell is inflated with depressed ovate whorl section and moderately involute with a n m o w , deep The round, steep umiblical wall curves in the highly arched venter. There isno sign of ornamentation on the last half-whorl, while the penultimate half-whorl has some faint lateral swellings and obsolescent broad ventral plications. The venter and a small part of the flank of the penultimate whorl, however, display evenly ,paced, stout, blunt secondaries and the outer umbilical shoulder bears distant nodes (c. 5 per half-whorl) which change apicad into bullate primary costae. A blunt constriction is present at about 95 mm diameter, at the end of the penultimate phragmocone whorl The test is thick. The septa1 suture is simple with broad shallow lobes of which the three to four umbilical lobes are somewhat oblique. The last several sutures are approximated. Remarks This species resembles the associated G. loupekinei more than any other known species of Gravesia but differs by the more compressed and involute whorls, by the weaker ornament, and by the slightly larger size. Since high whorl compression, tight coiling, and weak ornamentation are normally correlated in infraspecific variation (Westermann, 1969), this specimen may well be a variant or morph (foma) of G. loupekinei Measurements SPECIMENNO.

D

U

U/D%

H

W

HIW

Family Aspidoceratidae Zittel, 1895 Subfamily Aspidoceratinae Zittel, 1895 Genus Aspidoceras Zittel, 1868

Type Species Ammonires rogoznicensis Zeuschner, 1846. Aspidoceras cf. A . acanthicum (Oppel, 1863) 9 PI. 14, figs. la-c; 2a.b; 3a,b

+ ? 8'

Plate I . C1g5. 1-3 I

P / ~ ~ l l ~ ~ c 1ro{rp111r1 ertrs Benecke Changamue Shale: H > h n o r u m Zone. Louer Tithonian. Cndi\toned phragmocone. lateral (a.h) and apenural Icr \-ieu,s KNMI- MA?^. LOC. 8a. Neu Mro Pdnga Q u a m , nonh ot' Freretou n. hlombaaa area.

? Calliph~llorrrasber~acetlse(Catullo) Changamwe Shale. H!bonotum Zone: Louer Tithonian. Phragmocone u ~ constriction h and septa1 sutures. lateral view KNMI- MA?^. Lnc. 8c. New Mto Panga Q u a m . nonh of Freretown. Mombasa area.

o C . ~rrolr~~arrron (Boehml 3 C a l l i p / ~ ~ l l o c c raff. Changamwe Shale. Beckeri Zone; Upper Kimmeridgian. Incomplete phragmocone. lateral 2 5 . 8b. nonh of Freretown. Mornbasa (a) and apertural f b ) \.iews, sutures ( c ) K N M I - ~ ~Loc. area. Scale for all fieures equals 1 cm

Plate 1. figc. 1-3 I P!\.chopl~~lkocerns pr\.choicunz (Quenstedt) Changamwe Shale: BeckeriiH~tanotumZones. L'pper KimmeridgianiLower T i t h , ~ Phragmoconc and hody chamber. apertural ( a ) and lateral ( b ) views. ROM 34602. LO' shore of Port Tudor. 2.75 miles north-west of Chanfamwe Station and 2 miles nonh-,kc,l o f junction of track to Nguu Tatu with main coast road. 2 Lyrocerosfroosi DacquC Changamu,e Shale, Beckeri/Hytanotum Zones; Upper Kimmeridgian/Lower Tithonian. 28 Phragmocone with body chamber, inner whorls missing. lateral view K N M I - ~ ~Loc 15b. Changamwe Peninsula, Kipevu, northeast comer of Port Reitz. 3

Plero!vloceros montanum (Oppel)

Changamwe Shale, Hybonotum Zone; Lower Tithonian. External mould with protocct. . :. (a), and part of the internal mould of crushed body chamber, lateral view (b) K N M I - ~ ~ 2 9 . Loc. 8d. New Mto Panga Quarry, Mombasa area. Scale for all figures equals 1 cm

Plate 3. ftgs. 1-3 I

Turur?~ellicerusfT~rr~~rncl/icerus) cf. T . kuchhurrse (Waagen) Changamwe Shale. H)bonotum Zone: L o u r r Tithontan. Phrapmocone and bod! c h a n ~ h r u r l l preserved on one side. lateral ( a ) and ventral ( b ) views K N M I - ~ ~Loc. 3 0 . 8d. Neu ~ I I C , Panga Quarry, Mombasa area.

2 Turur?lelliceras fTura~nrllicerrr~) rruch\-r~orur?~ (Oppel)? Changamwe Shale, Becheri Zone: Upper Kimmeridgtan. Body chamber fragment. la; ( a ) and ventral (b) views KNMI-~431.Loc. 8h. New klto Panga Quarry. Mombasa a:. 3 Srreblrtes hub!ensis Sparh Changamwe Shale, Hybonotum Zone: Lower Tithonian. Complete specimen with partiall\ crushed body chamber, lateral (a,b), ventral (c), and apertural (dl views K N M I - ~ ~ 3 2 . 8d, Ken Mto Panga Quarry. Mombasa area. Scale for all figures equals 1 cm.

Plate 4, figs. 1-2 I Karrolicerus porri~~gcri (Sowerby) Changamwe Shale. Beckeri Zone: Upper Klmmendgian. Phragmocone w ~ t h h. chamber, lateral (a) and ventral (b) views K N M I - ~ ~k3 c7 . 8h. New Mto Panga Quarr!. Mombasa area. 2 Knrrolicerus aff. K. porrrrlgeri (Sow.) 9 Changamwe Shale. Hybonotum Zone: Lower Tithonian. Nearl) complete with paniall! crushed body chamber, lateral (a), apenural ( b ) , and ventral I c ) views K N M I - ~ ~Loc. 3 8 . 8c. New Mto Panga Quarry, Mombasa area. Scale for all figures equals 1 cm

Plate 5. figs. 1-2 1 Subdichoro~~rocera.? aff. S . sparsrplrcurum (Waagen) Changamwe Shale. Beckeri or basal Hyhonotum Zone: hounda? K~mmeridgianITithonian. Almost complete with apenure missing, lateral (a), apenural tb), and ventral ( c l views K N M I - ~ ~ 4 Loc. 1 . 8b. New hlto Panga Quarry, Mombasa area.

2 h'orrolicerus? cf. K . barhylocus (Waagen) $ Changamwe Shale. Beckeri Zone: Upper Kimmeridzian. Microconch, slightly damaged 9. with somewhat crushed bod) chamber. lateral (a) and apenural (b) views K N M I - ~ ~ 3 Loc. 8h, New Mro Panga Quarry. Momhasa area. Scale for all figures equals I cm

Plate 6. figs. 1-3 1-2

Pach~sphO~ctes beviehi (Fut~erer) changarnwe Shale, Hybonotum Zone; Lower Tithonian. 1 . Phragmocone with partial body chamber, lateral view K N M I - ~ ~ 2. 3 6Phragmocone . with one-third whorl bod! 3 5 . 8d, New chamber well preserved. apenural (a) and lateral (b) views K N M I - ~ ~ Loc. Mto Panga Quany, Mombasa area. 3 Subdichotomoceras(?) cf. S. sparsipliratum (Waagen) Changamwe Shale, Hybonotum Zone; Lower T~thonian.Phragmocone with one-qum?. whorl body chamber, lateral view KNMI-MAN.Loc. 8d. Neu Mto Panga Quarr! Mombasa area. Scale for all figures equals 1 cm

Plate 7. fipa 1-2 s ( ?S ). sporsiplicalufn (N'aagen) 1-2 S ~ r b d i c l ~ o ~ o ~ n o c e r acf. Changamwe Shale. H)bonotum Zone: Lower Tithoman. 1. Less than one-th~rdu hurl hod! chamber or entlrel) seprate and partiall! c r u h e d , lateral ( a ) and ventral ~ h \)l e a \ K N ! . I ~ - M A2.~Entirel) ~ ~ ~ . beptate and partially crushed. lateral view K S ~ I I - M ALO< ~~~~. a d . I to 3 km nonh of Frsretoan. Mombasa area. Scale for all figures equals I urn

Plate 8. figs. 1-3 Lirl~ococeras?rr~or~~bussirrrrr~~r (Dacque) Changamwe Shale, Hybonotum Zone; Lower Tithonian. Well-preserved juvenile?, latersl (a) and apertural (b) vieu.5 K N M I - ~ ~ 4Loc. 3 b . 8d. New Mto Panga Quarr). Mombasa area. 2 Lirhacoceras cf. L . fraasi (Dacqui) Changamwe Shale. Hybonotum Zone: Lower Tithonian. Partly crushed and incomplete phragmocone, lateral vieu K N M I - ~ ~k4 c2 . 8c. New Mto Panga Quarr). Mombasa are:! 3 Lirhacoceras cf. L. ulhullrrn (Quenstedt) Changamwe Shale. Beckeri Zone; Upper Kimmeridgian. Phra,omocons u ith one-li. whorl damaged body chamber w~rhoutaperture. lateral view KNMI- MA^. Loc. 8b. h e \ , Mto Panga Quarry. Mombasa area. 1

Scale for all figures equals 1 cm

Plate 10. figs. 1-3

Phanerosrephanus (Nothostephanus) digoi sp. nov. 0 , paratypes Changamwe Shale, Hybonotum Zone; Lower Tithonian... 1 Adult shell with one-quarter whorl incomplete body chamber; full lateral view (a) and pan of outer whorls removed to show ribbed venter of adolescent stage, lateral (b) and ventral (c) views KNMI-MAZlb. -

.

2

Isolated inner whorl exposing venter, lateral (a), apertural (b), and ventral (c) ,,ie, KNMI-MA45.

3

Phragmocone with pan of body chamber, inner whorls isolated, lateral (a) and ventral ( b , views K N M I - ~ ~Loc. 4 6 . 8e, New Mto Panga Quany, Mombasa area. Scale for all figures equals I cm.

Plate I I, figs. 1-3

1-2

Prorraspedires cf. P . ufricor~us(Zwierzicky) 9 Changamwe Shale, Upper Hybonotum Zone: Lower Tithonian. I. Phragmocone u ith pdnially preserved body chamber, lateral (a) and apenural (b) views ROM 34604. From 600 m nonhwest of New Mto Panga Shale Quarry (above bed 8e). 2. Incomplete 4 7 .Sf, New Mto Panga Quarry, Mombasa area. specimen, lateral \iew K N M I - ~ ~LOC.

3 Gro~,esiolorrpetirzei sp. n o v . 9 Paratype: Changamwe Shale, Hybonotum Zone; Lower Tithonian. Weathered phragmocone with body chamber, lateral views (a,b) K N M I - ~ ~LOC. 4 9 .8d, New Mto Panfa Quarry, Mombasa area. Scale for all fizures equals I cm

Plate 12, figs. 1-3 Gravesia loupekitlei sp. nov. 9 Changamwe Shale, Hybonotum Zone: Lower Tirhonian. 1 Holotype: complete slightly distorted internal mould, penultimate whorl damaged. lateral view (a) and penultimate whorl mostly removed to expose whorls. lateral (b) and apertural (c) views W M I - ~ ~ 2LOC. 2 . 8e. New Mto Panga Quany, Mombasa area. 2 Paratype: Incomplete phragmocone, lateral (a), apertural (b), and ventral (c) views R O ~ 34819. From about 3 km north-northwest of junction of track to Nguu Tatu and main coast road, Mombasa area. 3

Paratype: Outer whorl removed to expose the penultimate whorl. lateral view K N M I - ~ ~ 4 8 . Loc. 8d, New Mto Panga Quarry. Mombasa area. Scale for all figures equals 1 cm.

ate 13, fig. 1 Gra~,eziaaff. G. lorrpekiner sp. nov. 9 Changamwe Shale, H)bonotum Zone; Lower Tithonian. Well-preserved phragmocone. slightly damaged internal mould with test remains, complete, lateral (a) and ventral ( h l views, and, with part of outer whorl removed to expose ~ n n e rwhorl, lateral ( c ) and apertural (d) views KNMI-MASO. Loc. 8e. New Mto Panga Quarry, Mornbasa area.

Scale for all figures equals I cm.

Plate 14, figs. 1-3

+

I

Aspidoceras cf. A . acarlrhicum (Oppel) Q ? 8 Changamute Shale, Hybonotum Zone; Lower Tithonian. Phlagrnocone ( 9 ) with partial body chamber. lateral (a) and apenural (b) 5'lews, ' sutures (c) RoM 34820. From 500 to 600 m east of shore of Port Tudor and 4 km northeast of Changamwe Station.

2 Phragmocone ( Q ) with partial body chamber, lateral (a) and apenura] (b) views KNMI-MAS]. Loc. 8d. New Mto Panga Quany, Mombasa area. 3 Probable microconch, lateral (a) and apertural (b) vieujs K N M I - ~ ~ 5Loc. 2 . 8d. New Mto Panga Quarr)., Mombasa area. Scale for all figures equals 1 cm.

Plate 15, figs. 1-3 I

Aspidoceras aff. A. aca~irhicurn(Oppel) Changamwe Shale, Beckeri Zone: Upper Kirnmeridgian. Phragmocone, lateral view KNMI-MA%. LOC.8k, 1.3 km nonh of Freretown. Mombasa area.

2-3

Aspidoceras iphiceroide~Waagen 9 Changamwe Shale. Beckeri and Hybonotum Zones: Upper Kirnmeridgian and Lower 3. Tithonian. 2. Macroconch with missing inner whorls, lateral view K N M I - ~ ~ 7 7 3Loc. 8h, New Mto Panga Quany. Mombasa area. 3. Complete macroconch, lateral (a) and apenural (b) views ~ ~ ~ 1 . ~ ~ Loc 7 7 8d, 3 4New . Mto Panga Quarry, Mombasa area.

Scale for all figures equals I cm.

Plate 16. figs. 1-5

+6

1-3

Aspidoceras iphiceroides Waagen Q Changamwe Shale, Upper Kimmeridgian and Hybonotum Zone, Lower Tithonian. I. 5 6 . Ed, New Mto Complete microconch, lateral (a) and ventral (b) views K N M I - ~ ~LOC. PangaQuany, Mombasa area. 2. Spines, lateral view ~ ~ ~ 1 . ~ ~LOC. 7 7 8k, 3 5New . Mto Panga Quarry, Mombasa area. 3. Microconch with partial body chamber, lateral view KNMI-MA55. LOC.Ea, New Mto Panga Quarry, Mombasa area.

4-5

Aspidoceras cf. iphicerum (Oppel) Changamwe Shale, Beckeri and basal Hybonotum Zones; Upper KimmeridgianILower Tithonian. 4. Crushed internal mould with part of body chamber, lateral view KNMI-M~57a. LQC.Ea. New Mto Panga Quarry, Mombasa area. 5. Sutures K N M I - M A ~ ~ C . Loc. 8h, New Mto Panga Quarry, Mombasa area. Scale for all figures equals I cm.

Plate 17. figs. 1-3 I

Aspidoceras cf. A . appenninicurn Zittel

Changamwe Shale, Beckeri Zone; Upper Kimmeridgian. Panl) crushed large phragmocone without inner whorls. mostly internal mould, lateral views (a,b) K N M I - ~ ~ 5 LOC. 8. 8h, New Mto Panga Quarr). hlombasa area. 2 Aspidoceras (Ph~sodoceras)cf. A . a~~ellanurn Zittel Changamwe Shale. Hybonotum Zone: Lower Tithonian. Complete specimen with slightly . 8e, New damaged phragmocone, lateral (a) and apenural (b) views K N M I - ~ ~ 7 7 3 6LOC. Mto Panga Quarry, Mombasa area. 3

Aspidoceras (Ph~sodoceras)cicun~spinosrrm (Oppel)

Changamwe Shale, Hybonotum Zone; Lower Tithonian. Phragmocone with partially preserved umbilical spines, lateral (a) and apenural (b) views K N M I - M ~ 6 0 .Loc. 8e, New Mto Panga Q u a q , Mombasa area. Scale for all figures equals 1 cm.

.mJ I clenba 9ain%!j [[a JOJ a[eJS .earn eseqmow * h e n o e%ued oiw maN 'rg . m i ' ~ 1 9 v w - r w ~s m n a ! ~(q) renuan pue (e) piale[ '1uam3eq p o q 'ue!%p!iamm!y ~ i a d d n :auoz ! l a y ~ a g'aleqs amme%ueq3 ([addo) l I l ~ l ~ l l ~ q . .\ Hl / 'JJ S D ~ J J ! I O ~ I ~ ~Z. ~ H .earn eseqmow 'dsrena e3ued oity maN 'eg '207 . e [ g v w - r w ~ nsma!n (p) anbqqo pue '(3) [eiiuaA '(q) [elnl~ade'(e) p a l e l :JaqmaqJ Xpoq a q ~30 m d q11m a u o ~ o m % a ~ q.ue!uoql!~ d i a m g :auoZ mnloucqX~'aleqs ammeSueq3 ([addo) ~ n l 0 ~ 0 q . < $l ?{ J ~ J J ! ~ o u o ~ . < H 1

Plate 19, figs. 1-5

1 HJborroriceras pressulurn (Neumayr) Changamwe Shale, Beckeri Zone; Upper Kimmeridgian. Complete specimen. inner 3 a . 8h, New Mto Panga Quany, Mombasa whorls obscured, lateral view K N M I - ~ ~ 6Loc. area. 2 H~bonoricerash~bonorurn(Oppel) Changamwe Shale, Hybonotum Zone: Lower Tithonian. Part of the body chamber K N M I - ~ ~ 6LOC. l b . 8c, New Mto Panga Quarry, Mombasa area. 3 4 Hxborloriceras cf. H . onlarum (Spath) Changamwe Shale, basal Hybonotum Zone; Lower Tithonian. 3. Crushed fragment of body chamber internal mould K N M I - ~ ~ 6 24.a .Crushed fragment of body chamber b . from loc. 8a, New Mto PangaQuarry. Mombasa area. internal mould K N M I - ~ ~ 6 2Both 5 Hxbonoficeras cf. H . ciliarlrm Berckhemer and Holder Changamwe Shale, Hybonotum Zone; Lower Tithonian. Complete specimen with exposed inner whorls, lateral view KNMI-M~63b. Loc. 8c, New Mto Panga Quarry, Mombasa area. Scale for all figures equals 1 cm.

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1980

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WIMBLETON. W . A

1978

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