Early Cenomanian ammonites from East and North-East Greenland

Early Cenomanian (100.5–95.7 Ma) ammonite faunas from East and North-East Greenland collected by the late Simon Kelly and colleagues are described. The assemblages are dominated by typically boreal Schloenbachia varians (J. Sowerby 1817). Also present are Parapuzosia ( Austinic-eras ) austeni (Sharpe 1855) and species of more typically Tethyan genera. These include Phylloc-eras ( Hypophylloceras ) lombardense (Joly 2000), Gaudryceras ( Gaudryceras ) cassisianum (d’Orbigny 1850), Gaudryceras ( Mesogaudryceras ) leptonema (Sharpe 1855), and the hypermorphic tetrago-nitine Titanoleioceras boreale gen. et sp. nov. Previously known only from Geographical Society Ø and Traill Ø, the newly described material extends the distribution of these early Cenomanian faunas northwards to Hold with Hope and south to the Kangerlussuaq Basin. The phylloceratids, gaudryceratids, and tetragonitids in these assemblages are probably not preserved in their preferred original habitats, but rather drifted to their respective sites of burial during or after their lifetime.


Rationale and scope
In this contribution, we describe early Cenomanian (100.5-95.7 Ma) ammonites from East and North-East Greenland, most of which were collected by the late Simon Kelly during his time at CASP, UK.The presence of Cenomanian ammonite faunas in North-East Greenland was acknowledged by Leonard F. Spath (1946), who recognised the classic Boreal ammonite Schloenbachia amongst material collected by Hans Stauber on Geographical Society Ø and Traill Ø. Desmond Donovan, working in the same area, discovered additional material, documented and illustrated in a series of papers (Donovan 1949(Donovan , 1953(Donovan , 1954(Donovan , 1955;;summarised in Donovan 1957).The material described in this study extends the distribution of these faunas northwards to Hold with Hope and south to the Kangerlussuaq Basin (Kangersertuaq region; Fig. 1) and documents a greater diversity of taxa than previously recorded.
Simon R.A. Kelly passed away in May 2023, prior to the completion of this publication.Besides collecting most of the specimens, he had cleaned and prepared the material and made initial taxonomic and biostratigraphic assessments.Recognising the scientific value of the ammonites, he initiated the study and had seen and approved the photographic work and large parts of the text before his passing.As previously agreed, Simon Schneider joined as a co-author to complete Simon Kelly's tasks.W. James Kennedy is chiefly responsible for the taxonomic part of the study, and the new taxa take his authorship only.

Materials and locality details
The specimens described here were collected during several field seasons of CASP, between 1994 and 2003, and come from three regions in East and North-East Greenland (Figs 1,2).Farthest to the north, in the Hold with Hope region, Cenomanian ammonites were found in a single locality at Lygnaelv (CASP locality no.K7359, coordinates in decimal degrees 74.2639, -20.55, World Geodetic System 1984 (WGS84) datum; Supplementary File S1; 'HH' in Table 1), exposing the Fosdalen Formation (Home Forland Group; Kelly et al. 1998; see Bjerager et al. 2020 for updated lithostratigraphy).The Fosdalen Formation is an over 1-km thick unit of dark mudstones with siderite concretions, intercalated with varying proportions of thin fine-grained sandstone beds (Kelly et al. 1998), and was deposited in slope to basin floor settings according to Bjerager et al. (2020).Combined biostratigraphy of dinoflagellate cysts, inoceramid bivalves, and locally ammonites, indicates a middle Albian to middle Coniacian age (between 108 and 87 Ma) for the Fosdalen Formation on Hold with Hope (Kelly et al. 1998;Bjerager et al. 2020).Of three fragmentary Cenomanian ammonites collected, two are too poorly preserved for identification; the third specimen is assigned to Schloenbachia varians forma ventriosa below.

Kangerdlugssuaq Group
Kangerlussuaq Basin (Larsen et al. 2005  Approximately 100 km farther south, six localities on Geographical Society Ø also exposing the Fosdalen Formation, yielded ammonites (see Bjerager et al. 2020 for a revised lithostratigraphy).The Fosdalen Formation on Geographical Society Ø is thought to be of similar thickness as on Hold with Hope (cf.Bjerager et al. 2020) and represents slope to basin-floor deposits (Parsons et al. 2017).Note that Parsons et al. (2017, their supplementary materials 4) reported macrofauna including markers of early and late Albian, early and middle Cenomanian, late Turonian and Coniacian age from the Fosdalen Formation, supporting a chronostratigraphic range between 112 and 86 Ma.For the overlying Knudshoved Formation, sensu Bjerager et al. (2020), Parsons et al. (2017) reported late Santonian to early Campanian inoceramids and middle to late Campanian ammonites (indicating an age range between 84 and 70 Ma).Furthermore, Parsons et al. (2017) regarded the contact between the Fosdalen and Knudshoved Formations on Geographical Society Ø as conformable.In contrast, Bjerager et al. (2020) indicated a major hiatus encompassing the entire Coniacian and Santonian stages, but did not describe the contact of the two units on Geographical Society Ø. Thus, while the precise relationship between these two units remains unclear, the age restrictions imposed by Bjerager et al. (2020) are clearly contradicted by macrofossil evidence (Parsons et al. 2017).The lithostratigraphic column depicted in Fig. 2 represents a merger of both schemes.Cenomanian ammonites were collected from three localities in Tvaerdal (Fig. 1;  W4389,coordinates 72.8957,'LD2',CASP locality no. W4390,coordinates 72.8965,, and a single outcrop on Leitch Bjerg (Fig. 1; Table 1: 'LB', CASP locality no.W4342, coordinates 72.8555, -22.5023).All but one of these localities yielded specimens of Schloenbachia varians.In addition, Gaudryceras (Gaudryceras) cassisianum and Gaudryceras (Mesogaudryceras) leptonema each occur at a single locality (Table 1).
Another 500 km farther south, in the Kangerlussuaq Basin, ammonites occur in the Sorgenfri Formation, an up to c. 140 m thick unit of sandy mudstones with characteristic dark, phosphatic concretions, rare discrete sandstone beds, and occasional calcareous concretions and concretionary horizons (Larsen et al. 2005).Sedimentology suggests deposition in a mid-to outer shelf setting (Larsen et al. 2005).A middle Albian to middle Coniacian age (between 108 and 87 Ma) is indicated by combined ammonite and dinoflagellate cyst biostratigraphy (Larsen et al. 2005).Ammonites were collected from a single outcrop in 'Windy Valley' (CASP locality no. W4225,coordinates: 68.6383,, where the Sorgenfri Formation is more than 95 m thick (Fig. 3), consisting of mudstone with sparse bioturbation throughout, and concretions occuring in the lower 30 m of the succession.This locality has produced the richest and most diverse ammonite assemblage of Cenomanian age, including all six species described herein.Ammonites occur scattered within the lower 60 m of the exposure (Fig. 3), but most specimens were collected loose from the base of the outcrop.Based on the few specimens collected in situ, there is no apparent zonation.'Windy Valley' is the type locality of Titanoleioceras boreale gen.et sp.nov.
Generally, it should be noted that only the best-preserved, most representative specimens are included in this study.Numerous additional Cenomanian ammonite specimens collected by CASP are either too poorly preserved to be diagnostic or are more fragmentary representatives of the taxa described below (particularly Schloenbachia).

Discussion: age and composition of the faunas
The widespread occurrence of the genus Schloenbachia, identified as the early Cenomanian Schloenbachia varians (J.Sowerby 1817), confirms the horizon of the ammonites described below.This species dominates the overall assemblage and is a classic Boreal indicator, extending from North-East Greenland eastwards on the north side of Tethys to Iran north of the Zagros and western Kazakhstan (Fig. 4; Wilmsen & Mosavinia 2010, fig. 2;Kennedy 2013, text-fig. 1).Records from Algeria on the south side of Tethys (Kennedy & Juignet 1984, p. 123;Mendir et al. 2019, p. 239, plate 1, fig. 3) are based on poorly preserved Acompsoceras Hyatt 1903, in our view.Of other species present, Parapuzosia (Austiniceras) austeni (Sharpe 1855) was previously known from the Cenomanian and Turonian, ranging from southern England to France, Germany, the Czech Republic, Ukraine (Crimea), and, perhaps, KwaZulu-Natal in South Africa (Wright & Kennedy 1984, p. 60)  In contrast, cosmopolitan members of the Acanthoceratoidea, which provide zonal indices for the Cenomanian of the Eurasian part of the Boreal Realm, are absent, as are heteromorphs.
In lower Cenomanian successions, such as those in the West Melbury Formation marly chalks of south-east England or the phosphatic faunas of the Glauconitic Marl in the Isle of Wight, specimens of Schloenbachia varians make up perhaps as much as 99% of the ammonite fauna.Absence of Acanthoceratoidea and heteromorphs from the much smaller collections from East and North-East Greenland may thus simply reflect collection failure.The phylloceratid, gaudryceratid, and tetragonitid occurrences are unlikely to be a record of populations in their optimum habitats.Rather, they probably represented individuals that became separated from their parent populations, and drifted, either in life or after death, to their final site of burial.Quite where those parent populations were located is unknown.Dispersal of ammonites over long distances is documented and discussed by Kennedy & Cobban (1976, p. 64; see in particular text-fig.19).
A recent example of such an occurrence even more remarkable than the present ones is the single gaudryceratid recorded from the late Cenomanian and earliest Turonian hydrothermal vent faunas of the Troodos Massif in Cyprus that formed at depths of 2500-5000 m on arc-related spreading ridges of the Neotethys (Kaim et al. 2021(Kaim et al. , p. 1942, text-fig. 5f), text-fig. 5f), and interpreted as a waterlogged shell that sank from surface waters.

Systematic palaeontology
Conventions.Dimensions are given in mm: D: diameter.Wb: whorl breadth.Wh: whorl height.U: umbilicus.Figures given in parentheses are dimensions as a percentage of diameter.The suture terminology is that of Korn et al. (2003)  ?Discussion.Kennedy (1994)      Description.Phragmocones are up to 30.4 mm in diameter.Coiling is evolute, the broad umbilicus comprising up to 33% of the diameter, shallow, with a convex umbilical wall and shoulder.The whorl section is circular to slightly compressed (whorl breadth to height ratios down to 0.92).Internal moulds are smooth (MGUH 34487; Fig. 7I-K).Most specimens retain a recrystallised shell, the surface of which is ornamented by wiry primary ribs that are much narrower than the interspaces.
The ribs arise at the umbilical seam and strengthen across the umbilical wall (where they are markedly prorsiradiate).The ribs are prorsiradiate, strengthening progressively across the flanks, ventrolateral shoulders and venter, sinuous, feebly convex on ventrolateral shoulder and innermost flank, convex on the outer flank, sweeping forwards across the ventrolateral shoulder, and crossing the venter in a marked convexity.Fragments of body chamber associated with MGUH 34457 (Fig. 7F-H), together with GM 2024.2 (not figured), show this style of ornament extending to estimated whorl heights of up to 20 mm.In some specimens, occasional ribs are strengthened and form feeble flares (MGUH 34488: Fig. 7A-D).The septal lobe is very large (Fig. 7F, I).
Discussion.The present specimens differ in no significant respects from the holotype, and material from southeast France described by Thomel (1987Thomel ( , 1992)), Wright & Kennedy (1984), and Kennedy (1994)   Diagnosis.Giant Tetragonitinae with phragmocones up to 230 mm in diameter.Whorl section as wide as high or depressed, trapezoidal.Smooth but for growth lines and striae during early growth stages.Feeble, very widely separated constrictions appear in middle growth and are prominent on the adult body chamber; straight and prorsiradiate on the flanks, flexed back on ventrolateral shoulders and feebly concave over venter.Striations develop on the adult body chamber.Suture (Fig. 9) with irregularly trifid saddles and bifid lobes and small auxiliary saddles; septal lobe massive.Suture becomes highly subdivided at maturity.Discussion.Titanoleioceras is interpreted as a hypermorphic giant derivative of Tetragonites, reaching a similar size to the largest species of Pseudophyllites Kossmat 1895.It differs from that genus in the persistence of constrictions    Type locality and horizon.All specimens are from the CASP locality W4225, 'Windy Valley', Kangerlussuaq Basin (coordinates: 68.6383, -30.9297WGS84 datum), from the Sorgenfri Formation.Several specimens of Titanoleioceras boreale occur below and above a horizon that yielded Schloenbachia varians, which confirms their early Cenomanian age (Fig. 3).However, most specimens were collected from scree at the base of the section.

Diagnosis. With the characters of the genus.
Dimensions.Measurements on four paratypes (MGUH 34465,MGUH 34455,MGUH 34456,and MGUH 34454) GEUSBULLETIN.ORG      in diameter, with a single feeble constriction towards the adapertural end.The outer whorl of the holotype (Figs 10-12) is a 120° sector from the adapical end of the adult body chamber.Two prominent constrictions are present, one towards the adapical end, the other at the adapertural end.They are very feebly concave on the umbilical wall, prorsiradiate on the flanks, across which they broaden and deepen, flex back, and are feebly convex on the ventrolateral shoulder and cross the venter in a very feeble convexity.Growth lines and shallow grooves parallel the constrictions, and feeble strigations are present, most conspicuous on the ventrolateral shoulders and venter where the shell is preserved but effaced on the internal mould.
The suture is described in the diagnosis for the genus.Spath (1922, p. 127).Description.MGUH 34462 (Fig. 18B-D) may be a microconch phragmocone with an original estimated diameter of 100-110 mm, retaining traces of recrystallised shell.Coiling is evolute, with 48% of the previous whorl covered, the umbilicus broad, shallow, with a low, feebly convex wall, and narrowly rounded umbilical shoulder.The whorl section is compressed, with a whorl breadth to height ratio of 0.7.The greatest breadth is below mid-flank.The inner flanks are flattened, the middle and outer flanks feebly convex, converging to broadly arched ventrolateral shoulders and venter.There appear to be as many as eight constrictions on the inner flanks of the penultimate whorl.These are narrow, straight, and feebly prorsiradiate, with no other ornament on the surface of the intervening internal mould.On the outer whorl of the specimen, there are three constrictions per half whorl, deeply incised into the umbilical wall, strong and narrow on the flanks, straight and prorsiradiate on the inner flank, flexing back at mid-flank, feebly concave on the outer flank, sweeping forward across the ventrolateral shoulders, and crossing the venter in an obtuse linguoid peak.The adapical edge of the constrictions is strengthened into a feeble collar-rib.There are up to 16 ribs between successive constrictions.These parallel the constrictions, arising as mere striae on the inner flank, but strengthening on outer flanks, ventrolateral shoulders and venter, which bear an ornament of crowded, even, rounded ribs that are concave on outer flanks and ventrolateral shoulder and cross the venter in a rounded linguoid peak.At the adapertural end of the specimen, the last constriction and succeeding ribs modify their course, projecting strongly forward on the inner flank and back at mid-flank, to define the beginnings of lateral lappets (Fig. 18B).Traces of a further 240° of body chamber are preserved (Fig. 18C).MGUH 34461 (Fig. 17D, E) is a well-preserved phragmocone fragment with recrystallised shell preserved; the maximum preserved whorl height is 42 mm.The whorl breadth to height ratio is 0.74, the inner flanks are feebly convex and subparallel, the middle and outer flanks converging to the broadly rounded venter.Parts of two flared ribs are preserved, marking the adapical edge of constrictions on the internal mould.These collars are well-developed on the outer flanks, ventrolateral shoulders and venter, which they cross in an obtuse linguoid peak.

Parapuzosia (Austiniceras) austeni Sharpe 1855
Ornament is reduced to growth lines, striae, and riblets between the collars.= 48.7(30.4).The umbilicus is shallow, with a flattened, outward-inclined wall and very narrowly rounded umbilical shoulder.The flanks are flattened and subparallel, the ventrolateral shoulders and venter broadly rounded.There are two constrictions, and possibly a third at the adapical end.These are conspicuous on the internal mould, straight and feebly prorsiradiate on the inner flank, flexing back and concave at midflank, markedly concave on the outer flank, projecting forwards on the ventrolateral shoulders and effacing markedly on the venter, where they form an obtuse chevron.Ornament is very subdued on the internal mould.Where recrystallised shell is preserved, there is a strong adapical collar rib preserved, associated with the adapertural constriction.The shell surface between constrictions or collars is ornamented by more than 30 crowded, even ribs.These are straight, prorsiradiate, and weak on the inner flanks, flexing back and convex at mid-flank, strengthening and concave on the outer flank, and broadly convex and at their maximum strength over the venter.GM 2024.4 and GM 2024.7 (not figured) are small fragments of similar-sized individuals.MGUH 34460 (Figs 17A-C, 18A) is a further phragmocone fragment, with a maximum preserved whorl height of 110 mm.The whorl breadth to height ratio is 0.67.The umbilicus is shallow, the umbilical wall flattened and inclined outwards.The umbilical shoulder is very narrowly rounded, and sharply defined.The inner to middle flank region is feebly convex, the outer flanks convergent, the ventrolateral shoulders broadly rounded, the venter very feebly convex (Fig. 17B).The surface of the recrystallised shell is ornamented by crowded even ribs.These arise on the umbilical shoulder.They are straight and feebly prorsiradiate on the inner flank, flexed back and feebly convex at mid-flank, flexing back, and concave on the outer flanks.Additional short ribs intercalate on the outermost flanks, and all ribs strengthen and sweep forwards on the ventrolateral shoulders and cross the venter in a very broad convexity.
The suture (Figs 19A,20) is deeply and intricately subdivided, with a strongly retracted suspensive lobe.
Occurrence.Lower Cenomanian to middle Turonian.The geographic distribution extends from southern England to Germany, France, The Czech Republic, Ukraine (Crimea), East Greenland, and perhaps KwaZulu-Natal (South Africa).Description.The material is highly variable, as with populations from elsewhere.The formae recognised are, from strongly to weakly ornamented, ventriosa, varians, subtuberculata, intermedia, and subplana.Compressed individuals, such as MGUH 34484 (Fig. 21F,  G) and MGUH 34453 (Fig. 21A, B), have ribs that arise either singly or in pairs at the umbilical shoulder, with a very delicate umbilical tubercle and a slightly stronger inner lateral one, at which point the ribs commonly bifurcate, so that there are more ribs at the ventrolateral shoulder than at the umbilical shoulder.The ribs are straight and prorsiradiate on the innermost flank, but flex back and are feebly sinuous, feebly convex at mid-flank, and feebly concave on the outer flank.All ribs terminate in small ventral clavi on either side of the siphonal keel.These specimens correspond to the forma subvarians, although the presence of two tiny tubercles separates them from Sharpe's specimen (1853, plate 8, fig.7), in which respect they are transitional to forma subtuberculata.This form is well-represented by MGUH 34485 (Fig. 21C-E), a specimen 66 mm in diameter, with stronger umbilical and inner lateral tuberculation, the ribs showing a tendency to loop between lateral tubercles and ventral clavi.Other specimens of subvarians-subtuberculata type are GM 2024.8, GM 2024.10, and GM 2024.14 (not figured).GM 2024.11(not figured) is an external mould of a crushed individual 80 mm in diameter with stronger ornaments and thus transitional between subtuberculata and the holotype of varians.MGUH 34481 (Fig. 22L) is a further very crushed macroconch of this type.The ornament is very well-preserved.On the inner whorls there are 11-12 small umbilical bullae, each linked by a low rib to large subspinose inner lateral bullae.The outer whorl is preserved to a height of 45 mm.Coarser ribbed and tuberculate still are MGUH 34482 and MGUH 34490.The former (Fig. 21H, I) may be an adult microconch.The penultimate whorl bears weak umbilical and very strong, coarse lateral tubercles, as in passage forms between formae varians and ventriosa.The fragmentary body chamber, preserved to a whorl height of 36 mm, appears to show eccentric coiling, the umbilical wall is flat and outward-inclined.Small bullae perch on the umbilical shoulder and give rise to one or a pair of prorsiradiate ribs.Lateral tubercles have effaced.MGUH 34490 (Fig. 21J, K) is a half whorl of body chamber with a maximum preserved diameter of 120 mm.Coarse bullae perch on the umbilical shoulder and give rise to coarse prorsiradiate ribs.These link to strong inner lateral bullae that give rise to one or two ribs that sweep forward and link to coarse ventrolateral clavi.These decline towards the adapical end of the specimen and are replaced by narrow, sharp, strongly prorsiradiate ribs that efface before reaching the strong siphonal keel (Fig. 21K).This specimen appears to be a near-complete adult macroconch of a variant between formae varians and ventriosa.

Superfamily
Discussion.Specimens of Schloenbachia dominate the Cenomanian assemblages from East and North-East Greenland, as they do throughout the Eurasian part of the Boreal realm.The present material comprises typical variants of early Cenomanian varians following the interpretation of Kennedy in Wright & Kennedy (2015, p. 408 et seq.).Thus, three successive species are recognised: Schloenbachia varians (J.Sowerby 1817), which is restricted to the lower Cenomanian; Schloenbachia coupei (Brongniart 1822) of the lower middle Cenomanian Acanthoceras rhotomagense Zone, and Schloenbachia lymensis Spath 1926, which first appears in the upper middle Cenomanian Acanthoceras rhotomagense Zone and has its acme in the lower upper Cenomanian Calycoceras (Proeucalycoceras) guerangeri Zone.Differences between the species are set out in their diagnoses: Schloenbachia varians: "Adults range from 41 to at least 185 mm in adult diameter.The most strongly tuberculate individuals have lateral and ventrolateral tubercles only on most or all of the phragmocone; umbilical bullae are present on the adult body chamber.Compressed individuals in which ribs dominate over tubercles have umbilical and inner lateral tubercles, the latter linked to approximately twice as many ventrolateral clavi by a single rib; zigzag and looped ribbing is uncommon.Individuals with intermediate ornament may have outer flank ribs strengthened into weak or incipient bullae.Suture deeply incised, with bifid E/A and A/U 2 ; A trifid." (Kennedy in Wright & Kennedy 2015, p. 422).
Schloenbachia coupei (Brongniart 1822): "Adults range from 36 to 130 mm in diameter.All but the most feebly ornamented variants have umbilical, lateral and ventrolateral tubercles on the phragmocone and part or all of the body chamber, and a fourth row of outer lateral bullae may develop in near-adult and adult individuals.Suture with moderately incised bifid E/A and A/U 2 ; A trifid." (Kennedy in Wright & Kennedy 2015, p. 436).
Schloenbachia lymensis Spath 1926: "A small species; adults range from 27 to 66 mm in diameter.Strongly ornamented variants with umbilical and lateral bullae linked by a narrow bar-like rib on phragmocone, sometimes extending onto adapical end of adult body chamber, lateral bullae efface on body chamber, as do umbilical and ventrolateral clavi close to adult aperture.More compressed variants have straight, flexuous, falcoid or lautiform ribs and constrictions on the adult body chamber.The weakest ornamented variants lack lateral bullae on the phragmocone, the body chamber ornament reduced to delicate riblets and lirae, tiny ventrolatral clavi and strong constrictions.Suture […] only moderately incised, with broad bifid E/A and A/U 2 ; A narrow and trifid."(Kennedy in Wright & Kennedy 2015, p. 451).
Schloenbachia species show wide intraspecific variation (e.g.Wilmsen & Mosavinia 2010;Kennedy 2013), and previous authors introduced a host of specific and varietal names.Kaplan et al. (1998, p. 106) introduced the term forma as a non-Linnean term for morphological variants, and this approach, as developed by Kennedy (2013) and Kennedy in Wright & Kennedy (2015) is followed here.

Simon Kelly (16 October 1949 -19 May 2023): an appreciation
Simon joined the Cambridge Arctic Shelf Programme (now known by its acronym, CASP) as a consultant in 1984, conducting field-based research on Svalbard, prior to working as a palaeontologist with the British Antarctic Survey (1988)(1989)(1990)(1991)(1992)(1993)(1994).He returned to CASP in 1994, conducting fieldwork and integrated biostratigraphic studies in Azerbaijan, Kazakhstan, Arctic Canada and, most notably, East and North-East Greenland.Simon's principal research focused on the Jurassic and Cretaceous periods, specialising in molluscan biostratigraphy and palaeoecology.His research was especially influential in establishing stratigraphical schemes for East and North-East Greenland.Simon authored 70 scientific articles (see appendix in Schneider & Pointon 2023).Simon was also curator of CASP's geological collections until 2020, overseeing their transfer into a purpose-built rock store that now bears his name.In 2000, Simon was awarded the Polar Medal for his Arctic and Antarctic service.During the subsequent 19 years, he participated in a further 14 Arctic field seasons for which he was awarded a prestigious second clasp to his Polar Medal as part of the UK government's New Year Honours list 2023.
sinuous across the flanks and transverse over the venter.Two strong collars are present on the fragment.They follow the course of the riblets and striae and strengthen progressively across flanks, ventrolateral shoulders, and venter.

Table 1
Occurrences of ammonite species in the eight localities studied marked by '×'.
SubgenusPhylloceras (Hypophylloceras) Salfeld 1924 Type species.Phylloceras onoense Stanton 1895, p. 74, by monotypy.Fig.4SimplifiedearlyCenomanian(100.5-95.7 Ma) palaeogeography of Europe and adjacent regions.Some partial present-day coastlines are given for orientation where appropriate.Occurrences of Schloenbachia varians are marked by stars -those reported herein in red (1: Hold with Hope.2:GeographicalSocietyØ. 3: Kangerlussuaq Basin), previously published sites in green.Map modified from Wilmsen & Mosavinia (2010) and Kennedy (2013).Phylloceras (Hypophylloceras) lombardensis (Joly 2000) Figs 5A-D ?1954Phylloceras cf.velledae (Michelin); Donovan, p. 5. 2000 Hyporbulites lombardensis Joly, p. 163, plate 39, fig.11; text-fig.387.2009HyporbuliteslombardensisJoly2000;Klein et al. pp.90, 92.Type.The holotype, by original designation, is in the B. Joly Collection (presumably in private hands; no repository given) and is from the condensed lower and middle Cenomanian Banc des Lombards, Cassis, Bouches-du-Rhône, France.Material.MGUH 34489 and MGUH 34467 from CASP locality W4225, 'Windy Valley', Kangerlussuaq Basin.Description.MGUH 34489 (Figs 5A,B) is a 120° sector of phragmocone retaining traces of recrystallised shell, with a maximum preserved whorl height of 39 mm.Coiling is very involute, the tiny umbilicus with a convex, outward-inclined umbilical wall and rounded umbilical shoulder.The whorl section is compressed subrectangular, with a whorl breadth to height ratio of 0.64, the greatest breadth just outside the umbilical shoulder.The flanks are flattened and subparallel, converging ventrally.The ventrolateral shoulders are broadly rounded, the venter feebly convex.Delicate riblets arise as mere striae on the umbilical wall, and strengthen across the umbilical shoulder, where they are grouped in bundles.The riblets are very feebly convex across the umbilical shoulders, prorsiradiate and near straight on the flanks, flexing very feebly forwards to cross the venter in the feeblest of convexities.MGUH 34467 (Figs 5C, D) is a much larger internal mould fragment of the outermost flank, ventrolateral shoulder, and venter of a phragmocone.The riblets are very feebly prorsiradiate on the flanks and ventrolateral shoulder, and near-transverse over the venter.Discussion.These specimens are referred to P. (H.) lombardensis on the basis of whorl section and ornament, with the riblets tending to arise in bunches from the umbilical shoulder.Occurrence.Condensed lower or middle Cenomanian of Cassis, Bouches-du-Rhône, France, and East Greenland.