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Cainozoic Research, volume 15, number 1-2 (October 2015)
Cainozoic
Research
PUBLICATION OF THE ‘TERTIAIRY RESEARCH GROUP’
AND THE
‘WERKGROEP VOOR TERTIAIRE EN KWARTAIRE GEOLOGIE’
VOLUME 15, NUMBER 1-2 (October 2015)
CAINOZOIC RESEARCH
Volume 15, number 1-2 (October 2015)
Contents
Dr David Harrison, 1926 – 2015 - Taxonomist, Arabist, palaeontologist, medi-
cal practitioner, and founder of the Harrison Institute
Paul Bates & Malcolm Pearch
Dr Christopher King, December 1943 - January 2015, some personal recollec-
tions
David J. Ward
The Pyramidellidae (Mollusca: Gastropoda) from the Miocene Cantaure For-
mation of Venezuela
Bernard M. Landau & Patrick I. LaFollette
Eucyon sp. (Mammalia, Carnivora, Canidae, Caninae), an early dog from Mill-
Langenboom, The Netherlands
Noud Peters, Lorenzo Rook & John de Vos
The Pliocene locality Balgoy (province of Gelderland, The Netherlands) and a
new record of the great white shark, Carcharodon carcharias (Linnaeus, 1758)
Taco J. Bor & Werner J.M. Peters
The family Nassariidae (Gastropoda: Buccinoidea) from the late Neogene of
northwestern France
Frank Van Dingenen, Luc Ceulemans, Bernard M. Landau & Carlos Marques da Silva
Additions to the gastropod fauna of the Pliocene of Estepona, southwestern Spain,
3. The genus Plesiothyreus Cossmann, 1888 (Phenacolepadidae), with a note on its
presence in the Middle Miocene of Winterwijk, Miste, The Netherlands
Bernard M. Landau & André Jansen
Spaniodontella Andrussow in Goloubiatnikow, 1902 – a critical survey of use
and validity of the genus, and its relationship to the genus Alveinus Conrad,
1865 (Mollusca, Bivalvia: Glossoidea, Kelliellidae)
Arie W. Janssen, Ronald Janssen & Jaap van der Voort
New species of Neritidae (Gastropoda, Neritimorpha) from the Solent Group
(late Eocene and early Oligocene) of the Hampshire Basin
Malcolm Francis Symonds
Pourcy (Paris Basin, France): preliminary assessment of an early Eocene NW
European tropical coastal environment from molluscs and vertebrate fossils
E. Spijkerman, F.A.D. van Nieulande, F.P. Wesselingh, S. Reich & S. Tracey
ISSN 1570-0399
p. 3
p. 5
p. 13
p. 55
p. 59
p. 75
p. 123
p. 127
p. 147
p. 155
Cainozoic Research, 15(1-2), pp. 155-180, October 2015 155
Introduction
The classic locality of Pourcy is a small quarry locat-
ed to the west of the village of Pourcy (49°9’33.28”N
3°54’33.68”E) in lower parts of the valley of the river
Ardre on the southwestern side of the D386 road (Fig. 1).
The relatively rich fossil lumachelle there was rst noted
by M. Pistat and recorded by Tuniot (1902) who listed
53 mollusc species and also noted the presence of sh,
reptile and other vertebrate fossils. He gave a log of the
section from a sketch by Lucien Bellevoye, noting ve
distinct beds below the topsoil. Fritel (1910, p. 267) lat-
er included a rough version of this log in his geological
guide to the region and added two additional gastropod
Pourcy (Paris Basin, France): preliminary assessment of an
early Eocene NW European tropical coastal environment from
molluscs and vertebrate fossils
E. Spijkerman1,5, F.A.D. van Nieulande2, F.P. Wesselingh3, S. Reich3 & S. Tracey4 ,5
1 Zonnelaan 50, 1561 ES Krommenie, The Netherlands, E-mail: eddy.spijkerman@xs4all.nl
2 Scheldepoortstraat 56, 4339 BN Nieuw-en-St.-Joosland, The Netherlands; e-mail: frvannieul@zeelandnet.nl
3 Naturalis Biodiversity Center, P.O. box 9517, 2300 RA Leiden, The Netherlands; e-mail: frank.wesselingh@natu-
ralis.nl; sreich81@gmx.de
4 ICZN Secretariat, Natural History Museum, London SW7 5BD, England; e-mail: s.tracey@nhm.ac.uk
5 corresponding authors
Received 3 November 2014, revised version accepted 1 September 2015
We have studied the fossil fauna from the locality of Pourcy (Marne) in the northeast of the Paris Basin, France, housed in various
collections, in order to assess the nature and diversity of the fauna and gain an insight into the nature of the conditions that could
have produced this assemblage. The mollusc fauna is strongly indicative of varied tropical estuarine environments probably vegetated
with mangroves, but may also contain material derived from underlying mangrove facies. Both coastal marine and terrestrial mollusc
and vertebrate faunas are also represented. The terrestrial community contained indicators of subtropical rainforest lowland with
broad river banks and lakes, but some reworking of materials seems likely. The early-middle Ypresian Falun de Pourcy probably
reected ongoing estuarine/mangrove conditions that characterised the late Sparnacian period. Preliminary species lists of mollusc and
vertebrate fossils are given, together with a comparison of mollusc feeding guilds to those of a modern Indo-Pacic estuary system.
Key words: Ypresian, palaeoecology, biodiversity, mangrove, molluscs, vertebrates, NW Europe.
Résumé Français
Pourcy (Bassin de Paris, France, Marne): évaluation préliminaire d’un environnement tropical côtier nord-ouest européen de l’Éocène
inférieur d’après les mollusques et les vertébrés fossiles
D’après différentes collections, nous avons étudié la faune fossile du site de Pourcy (Marne) localisé au nord-est du bassin de Paris,
an de présenter la diversité de la faune et de reconstituer les conditions qui ont déterminé cet assemblage. La faune de mollusques
témoigne de différents environnements estuariens tropicaux probablement bordés de palétuviers mais pouvant aussi renfermer du
matériel provenant du faciès de mangroves. On note la présence de mollusques marins côtiers et terrestres et une faune de vertébrés.
L’association terrestre contient des indicateurs de forêts subtropicales de plaine avec des rivières à larges berges et des lacs uviaux.
Les faluns de l’Yprésien inférieur et moyen de Pourcy reètent probablement le développement des conditions estuariennes et de
mangroves qui ont caractérisé le Sparnacien supérieur. Une liste préliminaire d’espèces de mollusques et de vertébrés est donnée, et
les guildes alimentaires des mollusques sont comparées à ceux d’un système actuel estuarien de l’Indo-Pacique.
Mots-clés: Yprésien, paléoécologie, biodiversité, mangroves, mollusques, vertébrés, NW Europe.
156 Spijkerman et al. Pourcy: preliminary assessment of an early Eocene NW European tropical coastal environment
Figure 1. Location of the Pourcy localities: A. quarry, B. tem-
porary exposure at the farm Le Moulin de l’Ardre; early
Ypresian ‘Falun de Pourcy’ in green; unnamed late Ypres-
ian deposits in orange; Lutetian and later deposits in grey.
Figure 2. Averaged section in the quarry at Pourcy, based on a
sketch of the eastern face by L. Bellevoye in Tuniot (1902)
and various later eld notes.
0. Thin topsoil.
1. Buff cross-bedded ne sand. A shelly lenticle several cm
thick near the base 1-2 m.
2. Buff cross-bedded medium-ne sand, the upper part
largely decalcied with clay lenticles but containing ver-
tebrate fossils, the lower part packed with shells, mostly
mesohaline species, and containing <5 mm glauconitic
siliceous clasts, some resembling fragments of bioherms.
Beds 1 and 2 have an apparent dip northwards 3-4 m.
3. Unfossiliferous ne sand 0.5 m.
4. Clayey sand (‘terre sablonneuse) 0.5 m.
5. Clay seen to c. 0.5m. Beds 3, 4 and 5 horizontally bedded.
records. Leriche (1912) gave a photograph of the quarry
face indicating two distinct intervals and made some ob-
servations on the probable age of the deposit from the
known distribution of various molluscs. Based on these
accounts, together with eld notes by the present authors
and by J.J. Hooker (pers. comm.) we have made an esti-
mated log of the overall stratigraphy (Fig. 2) which was
seen to vary somewhat along the 16 m eastern face of the
quarry. The dominant feature of the section was a 3-4 m
‘falun’ or bed of sand, the lower part packed with shells
(bed 2), which appeared to be dipping northwards and
perhaps channeling into the underlying horizontally bed-
ded sands and clays (beds 3-5). The upper part of this
bed comprised largely decalcied sand with intercalated
clays containing sh teeth, otoliths and mammal remains
(A. Lawson pers. comm. See Appendix 2). Recent au-
thors (e.g. Duprat et al., 2010) have divided the falun into
‘lower levels’ placed in mammal zone MP8a and ‘upper
levels’ in MP9.
The quarry existed until the end of the 20th century, but is
now heavily overgrown and no longer accessible.
A temporary excavation in the eastern bank of the track
to the farm ‘Moulin de l’Ardre’ was later made a few hun-
dred metres to the west of the quarry, at 49°9’43.16”N
3°54’21.84”E (Jagt, 1982). This section showed approxi-
mately 2 m of homogenous yellow-brown fossiliferous
sand covered by 2 m of barren sand (pers. obs., ES 1990).
The fossil fauna was closely similar to that in the quarry
and the two sections were considered to be continuous.
The fossil-bearing levels comprise the ‘Falun de Pourcy’
which is considered to represent a late stage of the early
Eocene (basal Ypresian) Epernay Formation (Aubry et
al., 2005). Based on microinvertebrates, Smith & Smith
(2003) suggested a possible correlation with calcareous
nannoplankton zone NP10 which would make it consid-
erably older than the age of 53.27 ± 0.15 Ma calculated by
Cainozoic Research, 15(1-2), pp. 155-180, October 2015 157
Escarguel (1999, p. 269). Conicts in the dating of this
shell bed are discussed below.
Permission to visit these sites should be sought from the
landowners.
Geological setting
The Pourcy sites lie in the outer part of an Oligocene
outlier to the northeast between Pourcy and Chamery.
Overlying the Falun de Pourcy are later Ypresian (Cuis-
ian) sands and clayey sands, the line of contact running
close to the quarry, shortly to the south of the D386 as
shown on the Carte Geologique de la France (à 1/50.000,
BRGM, Fismes XXVII-12, 1987). These horizons were
not recognised in the sections studied, but 80 m to the
northeast of the quarry, a roadside exposure showed buff,
ne calcareous sands with the gastropods Bayania lactea
(Lamarck), Sigmesalia sp., Sycostoma sp. and Keilosto-
ma minus (Deshayes) within this mapped zone, but the
preservation indicated a Lutetian age . Within 1 km of
the quarry to the northeast, excavations have shown the
presence of middle Lutetian Calcaire Grossier Formation
overlain by late Lutetian sands. The areas mapped as late
Ypresian deposits were not specically identied in the
two sites investigated at Pourcy.
Material and methods
An inventory of the fossil collection of one of us (ES),
recovered from the Pourcy sites over the last 30 years,
has been supplemented with additional records from the
present authors and other workers and material in the col-
lections housed at Naturalis Biodiversity Center, Leiden,
The Netherlands. The individual fossils found at Pourcy
have been treated taxonomically in various publications
but no recent assessment of the whole fauna has been at-
tempted, which is one aim of this paper.
In mixed faunas such as that at Pourcy, containing eu-
haline, stenohaline/euryhaline as well as freshwater and
terrestrial species, with a predominance of taxa from
families today associated with tropical mangrove areas, it
is traditionally assumed that the palaeoenvironment was
a river estuary, thus explaining the means of transport of
the various faunal elements.
Following the publication of a comprehensive account
of the molluscs and ora of such a modern tropical river
estuary by Lozouet & Plaziat (2010) we thought it use-
ful to make an autecological assessment of the molluscs
of Pourcy from a direct comparison of the overall faunal
composition (including the feeding ecology) to investi-
gate whether Pourcy could be matched to one or more
subenvironments within a tropical estuarine system, and
whether possible reworking of earlier deposits should
be taken into consideration. We therefore compared the
fauna from both sites at Pourcy (combined) with sample
data from the Abatan (Bohol Island, Philippines) river
estuary/open-mouth bay transect from which seven grab
samples were assessed by Lozouet & Plaziat (2010). The
results of the comparison are given in Figs 108, 109.
The following collections from the quarry have been
studied for this paper: F.A.D. van Nieulande (Nieuw-en-
St.-Joosland, The Netherlands, now in Naturalis Biodi-
versity Center, Leiden), J.J. de Vos (Terneuzen, The Neth-
erlands; collection now in museum Het Warenhuis, Axel,
The Netherlands) and material from the department of
Fossil Mollusca of Naturalis Biodiversity Center (Leiden,
The Netherlands). The latter collections include material
donated by A. Haandrikman, H. van Haren, A.W. Jans-
sen, D. van der Mark and L. Staadt. From excavations at
Le Moulin de l’Ardre material was collected by E. Spij-
kerman (ES) of Krommenie, The Netherlands (including
ex coll. C. Deerenberg), J. van der Voort (Osterncappeln,
Germany) and from W. Groeneveld (now housed in the
Naturalis Biodiversity Center collections). To maximise
our coverage additional species in various private collec-
tions (A) have also been added to the lists where identi-
cations could be veried, although these specimens were
not included in the statistics.
Feeding guilds used in this study were herbivores/detriti-
vores (H), predatory carnivores (CP; including scavengers
and predators on foraminifers), parasites (P), suspension
feeders (SU), deposit feeders (D) and chemosymbiontic
deposit feeders (CD). Ecological information of molluscs
was derived from the Neogene Marine Biota of Tropical
America molluscan life habits database (Todd, 2001) and
the comprehensive ecological information provided by
Beesley et al. (1998).
Mollusc identications follow Cossmann & Pissarro
(1904-1906), Cossmann & Pissarro (1910-1913) and origi-
nal descriptions where necessary, with nomenclature
amended after Le Renard (1994, 1995), Le Renard &
Pacaud (1994, 1995), Pacaud & Le Renard (1995), Reid
et al. (2008), Pacaud & Harzhauser (2012) and Symonds
& Pacaud (2010). Fish teeth were identied using Casier
(1946), with reference to Leriche (1907) and Dutheil
(1991), using the updated systematics and nomenclature
of Cappetta & Nolf (2005) and Cappetta (2012). Turtle
remains were identied from De Broin (1977) and mam-
mal records and identications using Louis & Michaux
(1962), Michaux (1964, 1968), Russell et al. (1967), Rich
(1971), Hartenberger (1971), Sudré et al. (1983), Baudry
(1992), Hooker (1994, 1996, 2010), Escarguel (1999), Smith
& Smith (2003), Tabuce et al. (2006), Solé et al. (2011)
and Hand et al., 2015. Mammal systematics follow McK-
enna & Bell (1997).
One complication concerning the mammal faunas is that
apparently several taxa were present during the early
Eocene along the entire north coast of the Atlantic through
the Thule land bridge (McKenna, 1972; Godinot, 1981,
1982; Hooker, 1994;). The occurrence of species on both
American and European sides of the land bridge (Es-
carguel, 1999) may not be adequately reected in their
taxonomy (Simpson, 1929). For vertebrates recent taxo-
nomic revisions have led to considerable changes in the
characterization and number of taxa. Taxonomic revision
of American vertebrates is beyond the scope of this pa-
per. Therefore, the data presented here serve as a prelimi-
nary assessment of the faunal composition.
158 Spijkerman et al. Pourcy: preliminary assessment of an early Eocene NW European tropical coastal environment
Figures 3-12. Mollusca from the Epernay Formation of Pourcy.
Sizes: W = width, L = length, H = height, D = diameter. Suggested environmental indicators: T = terrestrial; FW = freshwater;
ME = normally associated with marine/open estuarine facies, although many of the species are also known from brackish facies of
the underlying Argiles à Lignites; [unmarked] = brackish, which is likely to include a continuum of lowered salinity palaeoenviron-
ments. RGM = Naturalis Biodiversity Center, Leiden. Most of the gured specimens are taken from private collections but will be
deposited with the Natural History Museum, London in due course.
3a-b. Barbatia modioliformis (Deshayes, 1829), right valve, W 26.4 mm (ME).
4. Trinacria inaequilateralis (d’Orbigny, 1850), interior of left valve, W 6.5 mm; exterior of right valve, W 6.5 mm (ME).
5a-b. Trigonodesma baudoni adelomorpha Cossmann, 1913, left valve, W 2.3 mm (ME).
6a-b. Brachidontes dutemplei (Deshayes, 1858) RGM.794332, right valve, W 8.5 mm (ME).
7a-b. Crassostrea cf. sparnacensis (Defrance in Deshayes, 1832), right (lower) valve, W 38 mm. Ostreid species at Pourcy are
frequently blackened and rolled and are not distinguishable on shell shape alone. This species somewhat resembles the gry-
phaeid Pycnodonte but lacks the vesicular shell structure that characterises that genus.
8. Crassostrea sparnacensis (Defrance in Deshayes, 1832), left (upper) valve exterior, W 25 x L 51.5 mm; typically elongate
form.
9a-b. Anomia casanovei Deshayes, 1858, left valve, W 17.8 mm (ME).
10. Palindonaia sp., fragmentary juvenile right valve, W 3 mm (FW).
11a-b. Saxolucina proxima sparnacensis (Deshayes, 1857), right valve, W 16 mm (ME).
12. Parvilucina nana (Deshayes, 1857), exterior of right valve, W 2 mm (ME).
Cainozoic Research, 15(1-2), pp. 155-180, October 2015 159
160 Spijkerman et al. Pourcy: preliminary assessment of an early Eocene NW European tropical coastal environment
Figures 13-25. Mollusca from the Epernay Formation of Pourcy.
For abbreviations see caption of Figs 3-12.
13a-b. Corbicula (s.lat.) cardioides (Deshayes, 1857), right valve, W 40.1 mm. A tumid smooth corbiculid intermediate between the
Thanetian C. (s.lat.) veneriformis (Deshayes) and the more angular C. (s.lat.) gravii (Deshayes), under which name it is often
recorded from Pourcy.
14. Corbicula (s.lat.) antiqua Férussac, 1822, right valve W 35 mm. As noted by Tuniot (1902) this appears to be a large but
stunted extreme morph of C. (s.lat.) cuneiformis as found at Pourcy.
15a-b. Corbicula (s.lat.) cuneiformis (Sowerby, 1817), RGM.794340, left valve, W 30.1 mm. Larger than the typical form from the
Woolwich Formation of England, and separated with difculty from C. (s.lat.) antiqua.
16a-b. Corbicula (s.lat.) arnoudii (Potiez & Michaud, 1839), left valve, W 10 mm.
17a-b. Tellinocyclas tellinoides (Férussac, 1822), right valve, W 17.2 mm. Much larger than the Thanetian T. angusta (Deshayes),
T. tellinoides appears to be restricted to a narrow chronostratigraphic interval and is useful for correlation, as noted below.
18. Eupera sublaevigata (d’Orbigny, 1850), adult right valve, W 3.3 mm and juvenile left valve, W 1.1 mm (FW).
19. Eupera denainvilliersi (de Boissy, 1848), interior of left valve, W 2.1 mm; exterior of left valve of another specimen, W 2.1
mm (FW).
20. Pisidium berellense (de Laubrière & Carez, 1880), incomplete left valve, W 2.1 mm (FW).
21. Pisidium cf. gosseleti (Leriche, 1899), interior of adult left valve, W 1.7 mm; exterior of juvenile right valve, W 1.2 mm (FW).
22. Sphenia terquemi Deshayes, 1857, right valve, W 5.1 mm (ME).
23. Sphenia acuta Staadt in Cossmann & Pissarro, 1913, left valve, W 2.3 mm (ME).
24a-b. Varicorbula arnouldii (Nyst, 1843), RGM.794336, right valve, W 6.9 mm (ME).
25. Teredo sp., interior of left valve, W 3.8 mm (ME).
Cainozoic Research, 15(1-2), pp. 155-180, October 2015 161
162 Spijkerman et al. Pourcy: preliminary assessment of an early Eocene NW European tropical coastal environment
Figures 26- 43. Mollusca from the Epernay Formation of Pourcy.
For abbreviations see caption of Figs 3-12.
26a-b. Neritodryas dutemplei (Deshayes, 1864), two views, H 15.8 mm.
27a-b. Neritodryas guillioui Symonds & Pacaud, 2010, two views, H 20 mm.
28a-b. Neritoplica uniplicata (Sowerby, 1823), two views, H 14 mm.
29. Neritoplica uniplicata (Sowerby, 1823), variety with colour banding, H 8.2 mm.
30a-b. Clithon barbei Symonds & Pacaud, 2010, two views, H 4.0 mm.
31a-b. Clithon sobrinum (Férussac, 1823), two views, H 9.9 mm.
32a-b. Clithon pisiforme pisiforme (Férussac, 1823), two views, H 4.5 mm.
33a-b. Clithon pisiforme perlongum (Cossmann & Pissarro, 1907), two views, H 6.5 mm.
34a-b. Viviparus rimatus (Michaud, 1837), two views, H 25.3 mm (FW). Recorded under various names, but there is no reason to
suppose that more than one species is represented.
35. Ampullina pistati Cossmann, 1907, H 42 mm (M).
36. Amaurellina lignitarum (Deshayes, 1864), H 6.1 mm (M).
37. Melanopsis antidiluviana (Poiret, 1797), H 20 mm. Abundant (see Discussion and conclusions).
38. Melanopsis cf. ovularis Watelet, 1853, H 7.2,mm. Rare.
39. Coptostylus pourcyensis (Cossmann, 1907), H 20.8 mm.
40. Coptostylus pourcyensis (Cossmann, 1907), apex of juvenile shell, enlarged (protoconch D 150 μm).
41. Coptostylus cf. albidus (Lamarck, 1804), H 10.4 mm. Pourcy examples have shallower sutures and smoother spires than typi-
cal C. albidus from the Sables de Cuise. This species lacks the denticles inside the lip and basal cords usually developed in
C. pourcyensis.
42. Hemisinus pistati (Cossmann, 1907), H 36 mm.
43. Bayania cf. hordacea (Lamarck, 1804 ), H 3.4 mm. Rare examples are somewhat waterworn (M).
Cainozoic Research, 15(1-2), pp. 155-180, October 2015 163
164 Spijkerman et al. Pourcy: preliminary assessment of an early Eocene NW European tropical coastal environment
Figures 44- 61. Mollusca from the Epernay Formation of Pourcy.
For abbreviations see caption of Figs 3-12.
44. Potamides turris (Deshayes, 1833), H 42 mm. Broad form resembling P. papalis (Lamarck, 1833) from the Sables de Cuise
but differing in the apical ontogeny. Narrower examples show considerable variation.
45. Potamides turris (Deshayes, 1833), H 37.5 mm. Conical morph with irregular projecting anges, that occurs throughout the
Epernay Formation.
46. Potamides funatus (Sowerby, 1814), H 40 mm. Very variable at Pourcy and includes forms approaching var. rillyensis (Coss-
mann) and var. cossmanni (Cossmann & Pissarro). May be hard to distinguish from P. turris and P. dimorphospira, and the
apical whorls, only present in juveniles, are often necessary to separate these species.
47. Potamides dimorphospira (Cossmann & Pissarro, 1913), H 27 mm. Characterised by the cyrtoconoid outline of the early
whorls.
48. Potamidopsis pourcyensis (Cossmann & Pissarro, 1913, H 11.9 mm. Uncommon and usually fragmentary at Pourcy, but
frequent in the Argiles à Lignites at Saran. The columella is swollen centrally, particularly in juveniles.
49. Vicinocerithium cf. biserialis (Deshayes, 1833), H 24.5 mm. Usually rolled and damaged at Pourcy but common in the Sables
de Cuise. Probably indicates a higher salinity than that of the potamidids.
50. Vicinocerithium cf. biserialis (Deshayes, 1833), juvenile, H 5.7 mm.
51. Vicinocerithium (s.lat.) scheri (Deshayes, 1864), H 48.2 mm. The broad tuberculate early whorls show that this does not
belong in the lineage of Vicinocerithium (s.str.).
52. Faunus cerithiformis (Watelet, 1855), H 47 mm. Also occurs in the Argiles à Lignites at Mutigny.
53. Faunus cerithiformis (Watelet, 1855), juvenile, H 10.1 mm, showing smooth attened early whorls.
54. Jponsia cuvieri (Deshayes, 1825), H 87 mm. The largest gastropod species at Pourcy, ranging up to the early Lutetian.
55. Jponsia cuvieri (Deshayes, 1825), juvenile, H 9.7 mm. Pacaud & Harzhauser (2012) described and gured a full growth-
series of this species.
56. Brotia praecessa (Deshayes, 1861), H 30.6 mm. Known also from late Thanetian strata.
57. Brotia melanioides melanioides (Sowerby, 1816), H 40 mm. Abundant and commonly waterworn. Protoconch suggests a
lecithotrophic larval life like many Potamides species, and probably a lagoonal habitat.
58. Brotia melanioides melanioides (Sowerby, 1816), apical whorls of young shell, H 2.8 mm. Juvenile whorls vary from convex
(as shown) to subcylindrical with attened sides.
59. Brotia melanioides bicoronata (Staadt in Cossmann, 1913), H 50 mm.
60. Haustator cf. comptus (Deshayes, 1861), juvenile, H 10 mm. Regarded as an indicator of shallow more euhaline conditions
than most of the fauna (ME).
61. Paludomus triticea (Férussac in Deshayes, 1825), H 2.8 mm.
Cainozoic Research, 15(1-2), pp. 155-180, October 2015 165
166 Spijkerman et al. Pourcy: preliminary assessment of an early Eocene NW European tropical coastal environment
Figures 62-83. Mollusca from the Epernay Formation of Pourcy.
For abbreviations see caption of Figs 3-12.
62. Hydrobia sparnacensis (Deshayes, 1862), damaged juvenile shell, H 1.5 mm.
63. Bythinella intermedia (Melleville,1843), H 1.9 mm.
64. Bythinella parkinsoni (Morris, 1854), H 2.0 mm. Somewhat variable in shape but always with deep sutures [syn. B. alta
(Deshayes)]. Also frequent in the slightly older Woolwich Formation in England.
65. Lapparentia elatior (Cossmann, 1907), H 3.8 mm. Also frequent in the slightly older Woolwich Formation in England
66. Lapparentia elatior (Cossmann, 1907), H 3.1 mm, with breakage exposing internal lamellae.
67. ‘Stenothyra’ chorista (Cossmann, 1888), H 1.7 mm.
68. ‘Stenothyra’ miliola (Melleville, 1840), H 1.4 mm.
69, 70. ‘Stenothyra’ cf. pulvis (Deshayes, 1861), two examples both H 1.4 mm. Apparently a form intermediate between ‘S’. miliola
and ‘S’. pulvis. While resembling the shape of some Stenothyridae, these Pourcy species lack the characteristic parietal ridge
in the aperture that holds the operculum in living stenothyrids, and these seem likely to be convergent hydrobiids.
71. ‘Stenothyra’ pulvis (Deshayes, 1861), H 1.3 mm.
72. Pasithea pourcyensis (Staadt in Cossmann & Pissarro, 1913), H 1.3 mm. Also frequent in the slightly older Woolwich Forma-
tion in England.
73. Cirsomphalus tunioti (Cossmann, 1902), H 2.4 mm. Introduced as Bithynia tunioti by Dollfus (in Tuniot, 1902) without
gure or description. The rst valid description was by Cossmann the same year.
74. Euspira consobrina (Deshayes, 1864), H 6.7 mm (ME).
75. Acirsa subtenuistriata moloti (Staadt in Cossmann & Pissarro, 1913), H 2.5 mm. Rare (ME).
76. Margineulima suturalis (Cossmann, 1907), H 2.5 mm. Rare (ME).
77. Cerithiopsis sp., H 2.8 mm. Rare (ME).
78. Cornulina praecursor (Cossmann, 1902),H 37.5 mm. Uncommon but probably over-represented in collections owing to bias.
Some living Melongenidae inhabit the littoral zone in brackish environments (ME).
79. Eocantharus latus (Sowerby, 1813), H 6.4 mm. Often associated with brackish molluscs in lagoonal facies in France and
England (ME).
80. Pseudoliva ssurata (Deshayes, 1835), H 8.3 mm. High-spired juveniles from Pourcy have been referred to P. laudunensis
despite closely resembling the Thanetian P. ssurata (e.g. Pacaud & Tracey, 2000, pl. 1, g. 5) (ME).
81. Pseudoliva laudunensis (Defrance, 1826), H 8.3 mm. This juvenile posesses the strong ribs and low spire that characterise
P. laudunensis. Further research should show any differences that could reliably separate the convergent P. ssurata (ME).
82. Eopleurotoma pourcyensis (Cossmann, 1902), H 9.4 mm (ME).
83. Domenginella ? sp., H 3.6 mm (ME).
Cainozoic Research, 15(1-2), pp. 155-180, October 2015 167
168 Spijkerman et al. Pourcy: preliminary assessment of an early Eocene NW European tropical coastal environment
Figures 84-100. Mollusca from the Epernay Formation of Pourcy.
For abbreviations see caption of Figs 3-12.
84. Anomalorbina hemistoma (Sowerby, 1816), left-apical view showing attened spire, D 1.3 mm; right-umbilical view of an-
other shell, D 1.3 mm.
85a-c. Anomalorbina cf. cuisensis (Cossmann, 1885), RGM.1008190, three views showing biconcave angular prole and strong
spiral ornament, D 1.0 mm. Usually with a few axial ribs on the early whorls. Arrangement of spiral threads varies and was
not mentioned in the original description of the type from the Sables de Cuise. Uncommon.
86. Valvata inexa Deshayes, 1862 ,apical view, D 1.6 mm (FW).
87. Odostomia lignitarum Deshayes, 1861, H 3.7 mm. Commonly associated with brackish molluscs in France and England
(ME).
88. Odostomia cf. primaeva Deshayes, 1861, H 2.2 mm (ME).
89. Odostomia sp., H 3.6 mm (ME).
90. Acteon sphaericulus granum (Cossmann, 1907), H 4.1 mm (ME).
91. Ringicula lignitarum Cossmann, 1902, H 2.8 mm (ME).
92. Carychium sparnacense (Deshayes, 1863), H. 1.5 mm. Modern Carychium live in wet situations, often near freshwater (T).
93. Carychium aff. cylindroides Staadt in Cossmann & Pissarro, 1913, H 1.3 mm. The original gures (Cossmann & Pissarro,
1913, g. 255-11) showed three shells which appear to represent two different species. The middle one of these most resem-
bles our g. 93 (T).
94. Carychium (s.l.) sp., H 1.4 mm. The size and half-ribbed ornament resembles some fossil species previously assigned to
Carychium, although the complex apertural dentition suggests afnities to certain Ellobiidae (T?).
95. Galba lignitarum (Deshayes, 1863), H 4.0 mm (FW).
96. Aplexa pulchella (d’Orbigny, 1850), H. 2.2 mm (FW).
97. Berellaia soluta Staadt in Cossmann & Pissarro, 1903, H 1.8 mm (FW).
98a-b. Biomphalaria subovata (Deshayes, 1825), RGM.794333, two views, D 4.3 mm (FW).
99. Acroloxus matheroni lemoinei (Cossmann, 1889), D 1.4 mm (FW).
100. Palaeostoa exarata (Michaud, 1838), ? juvenile, H 1.7 mm. Ribbed fragments of adult P. exarata are also found at Pourcy.
The fragility of many of the smaller freshwater and terrestrial species in the shell bed suggests that they were preserved in
the interiors of larger gastropods (T).
Cainozoic Research, 15(1-2), pp. 155-180, October 2015 169
170 Spijker man et al. Pourcy: preliminary assessment of an early Eocene N W European tropical coastal environment
Figures 101-107. Some characteristic vertebrate taxa from Pourcy (coll. ES, except Trionyx sp.).
101. Striatolamia macrota (Agassiz, 1843); lower anterior tooth, lingual view, H 19 mm.
102. Egertonia isodonta Cocchi, 1866; pharyngeal plate, oral view, H 22 mm.
103-104. Lepisosteus suessionensis Gervais, 1852; external view of scales, 103: H 11 mm; 104: H 11 mm.
105. Platychoerops daubrei (Lemoine, 1879); incisor tooth, labial view, H 19 mm.
106. Trionyx sp.; RGM n.n.; pleural plate of carapax, dorsal view, W 30.6 mm.
107. Gastornis parisiensis Hebert, 1855; humerus, L 60 mm.
Statistical results
The composition of the mollusc and vertebrate faunas are
provided in Tables 1 and 2. The mollusc fauna consisted
of 30 bivalve and about 80 gastropod species (Table 1).
The relative abundance of the common species in the
studied collections, veried by the cursorary examination
of an unsorted residue sample, was considered to provide
an adequate reection of actual frequency, although the
percentages of rarer taxa were likely to be exaggerated
owing to collecting bias. The bivalve fauna was domi-
nated by Corbiculidae (57%) and Corbulidae (24%), the
gastropod fauna by Neritidae (20%), Mela nopsidae (18%)
and Potamididae (18%). In mollusc species numbers,
slightly more than half of the Pourcy fauna was dominat-
ed by herbivores/detritivores. Suspension feeders made
up about one third and the combined groups of preda-
tors and parasites 14%. In abundance, suspension feeders
(mostly bivalves) made up almost half; carnivores were
rare (4%).
Cainozoic Research, 15(1-2), pp. 155-180, October 2015 171
28,29 Neritoplica uniplicata (Sowerby, 1823) 88 175
30 Clithon barbei Symonds & Pacaud, 2010 - A
31 Clithon sobrinum (Férussac, 1823) 251 236
32 Clithon pisiforme pisiforme (Férussac, 1823) - 19
33 Clithon pisiforme perlongum (Cossmann &
Pissarro, 1907) 32 11
Family Viviparidae
34 Viviparus rimatus (Michaud, 1837) 2 -
Family Ampullinidae
35 Ampullina pistati Cossmann, 1907 10 20
36 Amaurellina lignitarum (Deshayes, 1864) 44 36
Family Potamididae
44,45 Potamides turris (Deshayes, 1833) 246 237
46 Potamides funatus (Sowerby, 1814) 112 101
47 Potamides dimorphospira (Cossmann &
Pissarro, 1913) (b) - A
48 Potamidopsis pourcyensis (Cossmann &
Pissarro, 1913) 62 39
Family Batillariidae
49,50 Vicinocerithium cf. biserialis (Deshayes, 1833) - A
51 Vicinocerithium (s.l.) scheri (Deshayes, 1864) 27 5
Family Pachychilidae
57,58 Brotia melanioides melanioides (Sowerby,
1816) 150 143
59 Brotia melanioides bicoronata (Staadt in
Cossmann, 1913) 23 25
56 Brotia praecessa (Deshayes, 1861) 1 -
54,55 Jponsia cuvieri (Deshayes, 1825) 44 24
53 Faunus cerithiformis (Watelet, 1855) 27 23
Family Melanopsidae
37 Melanopsis antidiluviana (Poiret, 1797) 325 428
38 Melanopsis cf. ovularis Watelet, 1853 - A
39,40 Coptostylus pourcyensis (Cossmann, 1907) 40 36
41 Coptostylus cf. albidus (Lamarck, 1804) 5 -
Family Thiaridae
42 Hemisinus pistati (Cossmann, 1907) 8 3
Family Pseudomelaniidae
43 Bayania cf. hordacea (Lamarck, 1804 ) - A
Family Paludomidae
61 Paludomus triticea (Férussac in
Deshayes, 1825) 10 -
Family Turritellidae
Haustator cf. circumdatus (Deshayes, 1861)(x) 28 -
60 Haustator cf. comptus (Deshayes, 1861) 14 5
Ispharina cf. hybrida (Deshayes, 1835) (x) - 2
Family Hydrobiidae (s.l.)
62 Hydrobia sparnacensis (Deshayes, 1862) 208 64
Peringia glandinensis (De Laubrière &
Carez, 1880) (x) 1 2
Peringia sp. (x) 7 2
Pseudamnicola pistati (Cossmann, 1907) (x) 1 -
63 Bythinella intermedia (Melleville,1843) (x) - 19
64 Bythinella parkinsoni (Morris, 1854) 22 53
65,66 Lapparentia elatior (Cossmann, 1907) 16 3
Lapparentia stenochora (Cossmann, 1888)(x) 6 7
Staliopsis bouryi (Cossmann, 1888) (x) 23 1
73 Cirsomphalus tunioti (Cossmann, 1902) 17 4
67 ‘Stenothyra’ aff. chorista (Cossmann, 1888) 76 11
68 ‘Stenothyra’ aff. miliola (Melleville, 1840) - 14
69-71 ‘Stenothyra’ aff. pulvis (Deshayes, 1861) 14 1
Paladilhia plicistria (Cossmann, 1888) A -
Class Bivalvia
Family Arcidae
3 Barbatia modioliformis (Deshayes, 1829) 6 7
Family Noetiidae
4 Trinacria inaequilateralis (d’Orbigny, 1850) 61 23
5 Trigonodesma baudoni adelomorpha
Cossmann, 1913 7 5
Family Mytilidae
6 Brachidontes dutemplei (Deshayes, 1858) 120 19
Family Ostreidae
Ostrea cf. bellovacina (Lamarck, 1806) 15 12
7,8 Crassostrea sparnacensis (Defrance in
Deshayes, 1832) 86 155
Family Anomiidae
9 Anomia casanovei Deshayes, 1858 12 75
Family Unionidae
10 Palindonaia sp. - A
Family Lucinidae
11 Saxolucina proxima sparnacensis (Deshayes,
1857) 25 17
12 Parvilucina nana (Deshayes, 1857) - A
Family Leptonidae
Anomalokellia pourcyensis (Cossmann &
Pissarro, 1906) (a) - -
Family Cardiidae
Orthocardium moloti (Staadt in Cossmann &
Pissarro, 1913) (a) - A
Family Psammobiidae
Gari staadti (Cossmann & Pissarro, 1913) - 2
Family Corbiculidae
13 Corbicula (s.l.) cardioides (Deshayes, 1857) 31 24
14 Corbicula (s.l.) antiqua Férussac, 1822 314 876
15 Corbicula (s.l.) arnoudii (Pot. & Mich., 1839) 226 96
16 Corbicula (s.l.) cuneiformis (Sowerby, 1817) 100 230
17 Tellinocyclas tellinoides (Férussac, 1822) 32 13
Donacopsis heberti (Deshayes, 1857)(x) 1 -
Familie Sphaeriidae
20 Pisidium berellense (de Laubrière & Carez,
1880) - 1
21 Pisidium cf. gosseleti (Leriche, 1899) - A
18 Eupera sublaevigata (d’Orbigny, 1850) - A
19 Eupera denainvilliersi (de Boissy, 1848) - A
Familie Veneridae
Pitar lamberti (Deshayes, 1857) 18 1
Family Myidae
23 Sphenia acuta Staadt in Cossmann &
Pissarro, 1913 7 -
22 Sphenia terquemi Deshayes, 1857 11 6
Family Corbulidae
24 Varicorbula arnouldii (Nyst, 1843) 435 420
Family Pholadidae
Cyrtopleura orbignyana (Lévesque in
Graves, 1847)(a) - A
Teredina personata Lamarck, 1806 5 8
25 Teredo sp. - A
Class Gastropoda
Family Neritidae
26 Neritodryas dutemplei (Deshayes, 1864) 27 4
27 Neritodryas guillioui Symonds & Pacaud,
2010 - 1
Q M Q M
Table 1, continued next page
172 Spijkerman et al. Pourcy: preliminary assessment of an early Eocene NW European tropical coastal environment
gin of various lithologies resulting from deltaic/estuarine
environments. The closest match to the Falun de Pourcy
was ‘Medium- to ne-grained sandstones with bioclasts
and small mudclasts. Cross-bedding, normal grading;
traction carpets, HCS. Large foraminifera, mollusks.
Densely packed and well-sorted beds and pavements.
[interpreted as] Transported (sedimentologic) distal delta
front (shelfal lobes)’. Large foraminifera were, however,
not common at Pourcy.
Vegetation
Following Ellison et al. (1999), Plaziat et al. (2001) ex-
amined the history of mangroves in detail and concluded
they had existed at least since earliest Paleocene times,
and that the early Tertiary was a period of pan-tropical/
subtropical homogeneity of the mangrove ecosystem.
Pollen and spores of the extant mangrove plants Nypa,
Bruguiera and Acrostichum aureum have been found in
samples of the uppermost Argiles à Lignites d’Epernay
The vertebrate fauna consisted of 4 shark, 4 batoid, 7
bony sh, 5 turtle, 2 crocodilian, 1 bird and 34 mammal
species (Table 2). Some characteristic species are shown
in Figs 101-107.
The shark and batoid fauna was dominated by Odontas-
pididae/Mitsukurinidae (46%; the shark teeth were pre-
dominately of juvenile animals) and Myliobatidae (46%).
The bony sh fauna (represented by teeth and scales)
was dominated by Lepisosteidae (47%) and Phylodontidae
(29%). Otoliths were rather rare and those recorded are
listed in Appendix 2. The reptiles comprised crocodil-
ians (50%) and turtles (50%). Only a few remains of birds
and mammals have been recorded.
Discussion and conclusions
Sedimentation
Harzhauser et al. (2015) gave a simplied estimate of ori-
Family Iravadiidae
72 Pasithea pourcyensis (Staadt in Cossmann &
Pissarro, 1913)(b) - A
Family Naticidae
74 Euspira consobrina (Deshayes, 1864) 42 71
Family Epitoniidae
75 Acirsa subtenuistriata moloti (Staadt in
Cossmann & Pissarro, 1913)(b) - A
Cirsotrema stueri (De Boury, 1890) (x) 1 1
Family Eulimidae
76 Margineulima suturalis (Cossmann, 1907)(b) - A
Family Cerithiopsidae
77 Cerithiopsis sp. - A
Family Muricidae
Nucellopsis sarroniensis (Carez, 1879) - A
Family Turbinellidae
78 Cornulina praecursor (Cossmann, 1902) 2 9
Family Buccinidae
79 Eocantharus latus (Sowerby, 1813) 18 35
Family Pseudolividae
80 Pseudoliva ssurata (Deshayes, 1835) - A
81 Pseudoliva laudunensis (Defrance, 1826) 15 6
Family Turridae
82 Eopleurotoma pourcyensis (Cossmann, 1902) 5 5
Family Borsoniidae?
83 Domenginella ? sp. - A
Family Cornirostridae
84 Anomalorbina hemistoma (Sowerby, 1816) 61 101
85 Anomalorbina cf. cuisensis (Cossmann, 1885) 1 A
Family Valvatidae
86 Valvata inexa Deshayes, 1862 - A
Family Pyramidellidae
88 Odostomia primaeva Deshayes, 1861(x) 1 -
Odostomia asthenoptyxis (Cossmann, 1907) (b) - -
87 Odostomia lignitarum Deshayes, 1861 44 25
Odostomia gravesi Deshayes, 1861(x) 8 4
Odostomia microscopica (Cossmann, 1907) - A
89 Odostomia sp. - A
Syrnola sp. 1 1
Family Acteonidae
90 Acteon sphaericulus granum (Cossmann, 1907)(b)
- A
Crenilabium pourcyense (Cossmann, 1907) (b) - -
Family Ringiculidae
91 Ringicula lignitarum Cossmann, 1902 3 -
Family Retusidae
Retusa lignitarum (Cossmann, 1907) 2 -
Family Ellobiidae
Stolidoma pistati Cossmann, 1907(b) - -
Family Carychiidae
92 Carychium sparnacense (Deshayes, 1863) 1 -
93 Carychium cylindroides Staadt in Cossmann &
Pissarro, 1913(b) 1 -
94 Carychium sp. - A
Family Lymnaeidae
95 Galba lignitarum (Deshayes,1863) - A
Family Physidae
96 Aplexa pulchella (d’Orbigny, 1850) 1 -
Berellaia bonneti Cossmann, 1907(b) - -
97 Berellaia soluta Staadt in Cossmann &
Pissarro, 1903(b) 1 A
Family Planorbidae
98 Biomphalaria subovata (Deshayes, 1825) 23 10
Family Acroloxidae
99 Acroloxus matheroni lemoinei (Cossmann, 1889) - A
Family Megaspiridae
100 Palaeostoa exarata (Michaud, 1838) - A
Q M Q M
Table 1, continued
Table 1. Molluscs from Pourcy. For bivalves the number of valves is indicated. First column (Q): Quarry. Second column (M):
outcrop ‘Le Moulin de l’Ardre’. Records from the literature: (a) Cossmann & Pissarro (1906); (b) Cossmann & Pissarro (1913); (c)
Symonds & Pacaud (2010); (x) (record not veried). A= additional material noted in private collections. Bold numbers in front of
species refer to gures.
Cainozoic Research, 15(1-2), pp. 155-180, October 2015 173
Phylum Chordata
Class Chondrichthyes
Family Mitsukurinidae
Striatolamia macrota (Agassiz, 1843)(a,b,e) - 38
Family Odontaspididae
Brachycarcharias lerichei (Casier, 1946)(a,b,e) - 14
Hypotodus verticalis (Agassiz, 1843)(a,b,e) - 40
Family Carcharhinidae
Physogaleus secundus (Winkler, 1876)(a,b) - 14
Family Pristidae
Pristis sp.(a) - -
Family Myliobatidae
Myliobatis sp.(a) - 92
Aetobatus irregularis Agassiz, 1843(f) - -
Family Mobulidae
Burnhamia daviesi (Woodward, 1889) (b) - -
Class Osteichthyes
Family Lepisosteidae
Lepisosteus suessionensis Gervais, 1852(a) - 49
(29 scales, 20 teeth)
Family Amiidae
Amia (Pappicthys) barroisi Leriche, 1905(a) - -
Family Phylodontidae
Egertonia isodonta Cocchi, 1866(a) - 29
(7 jaws, 1 scale, 21 teeth)
Phyllodus toliapicus Agassiz, 1844(a,f) - 1
(1 tooth)
Family Ariidae
Arius egertoni Dixon, 1850 - 14
(1 jaw, 1 scale, 12 spines)
Family Sparidae
Sparidae indet. (otoliths) 1 1
Family Labridae
Labrodon trapezoidalis Leriche, 1905(a) (jaws) - 9
Class Reptilia
Family Petomedusidae
Neochelys sp. (c) - -
Family Trionychidae
Palaeotrionyx vittatus (Pomel, 1847) (c) - 3
Trionyx sp. (c) - 32
Family Carettochelyidae
Allaeochelys sp. (c) - 3
Family Testudinidae
Palaeochelys sp. (c) - -
Family Crocodylidae
Diplocynodon sp. - 36
Family Alligatoridae
Allognathosuchus sp. - 2
Class Aves
Family Gastornithidae
Gastornis parisiensis Hebert, 1855 - 2
Class Mammalia
Family Herpetotheriidae
Amphiperatherium maximum Crochet, 1979(d) - -
Peratherium sp.(g) - -
Family Paramyidae
Ailuravus michauxi Hartenberger, 1975(h) - -
Euromys thaleri (Michaux, 1964)(h,i) - -
Pseudoparamys teilhardi (Wood, 1962)(h,i) - -
Pantrogna russelli (Michaux, 1964) (h,i,j) - -
‘Paramys’ woodi Michaux, 1964(h) - -
Sparnacomys chandoni (Hartenberger, 1971)(d,k) - -
Family Apatemyidae
Apatemys sigogneaui Russell et al., 1979(d,m) - -
Apatemys mutiniacus Russell et al, 1979(d) - -
Family Esthonychidae
Plesiesthonyx minimus (Baudry, 1992)(n)(t) - -
Plesiesthonyx luciae (Baudry, 1992)(n)(t) - -
Family Coryphodontidae
Coryphodon (?) eocaenus Owen, 1846(d,g) - -
Family Paroxyclaenidae
Paroxyclaenidae indet.(o) - -
Family Oxyaenidae
Palaeonictis gigantea de Blainville, 1842(p) - -
Family Amphilemuridae
Placentidens latus Russell et al., 1973(d) - -
Neomatronella luciannae (Russell et al.1975) (d) - -
Family Onychonycteridae
Marnenycteris michauxi Hand et al., 2015 (f) - 1
Family Plesiadapidae
Platychoerops daubrei (Lemoine, 1879) (d,l) - 1
Plesiadapis cf. remensis Lemoine, 1887(g,l) - -
Family Paromomyidae
Phenacolemur fuscus (Russell et al, 1967) (d) - 1
Family Adapidae
Protoadapis curvicuspidens (Lemoine, 1878)(g) - 1
Cantius eppsi (Cooper, 1932) (d,l) - 1
Family Arctocyonidae
Landenodon woutersi Quinet, 1966(d,h) - 3
Family Louisinidae(u)
Paschatherium dolloi (Teilhard de Chardin,
1927) (d) - 2
Teilhardimys musculus (Teilhard de Chardin,
1927) (d,g) - -
Family Phenacodontidae
Phenacodus teilhardi Simpson, 1929(g) - -
Family Dichobunidae
Diacodexis varleti Sudré et al., 1983(d,q) - -
Protodichobune oweni Lemoine, 1878(g) - -
Bunophorus cappetai Sudré et al., 1983(d,q) - -
Family uncertain
Hallensia louisi Hooker, 1994(r) - -
Superfamily Equoidea
Pliolophus vulpiceps Owen, 1858(d,r,s) - -
Hyracotherium aff. leporinum Owen, 1841(d,g,r) - 1
‘Propachynolophus’ sp.(r) - -
Q M Q M
Table 2. Vertebrate fossils from Pourcy. First row (Q): Quarry. Second row (M): Outcrop ‘Le Moulin de l’Ardre’. References: (a)
Leriche, 1907; (b)Dutheil, 1991; (c)De Broin, 1977; (d)Hooker, 1996; (e)Casier, 1946; (f)Hand et al., 2015; (g) Louis et al., 1962; (h)
Escarguel, 1999; (i)Michaux, 1964; (j)Michaux, 1968; (k)Hartenberger, 1971; (l)Russell et al., 1967; (m, n))Baudry, 1992; (o)Rich et al.,
1971; (p)Solé et al., 2011; (q)Sudré et al., 1983; (r)Hooker, 1994; (s)Smith & Smith, 2003; (t)Hooker, 2010 ; (u)Hooker & Russell, 2012.
174 Spijkerman et al. Pourcy: preliminary assessment of an early Eocene NW European tropical coastal environment
at Verzenay, about 16 km east of Pourcy and at localities
near Epernay, southeast of Pourcy (Gruas-Cavagnetto et
al., 1980a, 1980b). From these, and from the lithology at
Verzenay, the authors deduced the existence of a man-
grove soil in this interval. Although not proven, the Fa-
lun de Pourcy must be either laterally equivalent to this
palaeosol or later in date, perhaps channeling into pene-
contemporaneous beds. Mangroves today are restricted
to the tropical/subtropical regions of the world and their
presence in the Epernay Formation suggests similar cli-
matic conditions at the time of the Pourcy deposits.
Dominant molluscs
The most abundant molluscs in the Pourcy fauna com-
prised Corbiculidae, Corbulidae, Neritidae, Melanopsi-
dae, Potamididae and Hydrobiidae.
Corbiculidae1
Among the extant Corbiculidae the genus Corbicula (s.s.)
is characterised by a thick, rounded equilateral shell with
strong concentric folds. It lives in rivers and is restricted
to freshwater, tolerating only the lowest salinity of tidal
reaches, however this genus is not represented in the early
Cenozoic. With the exception of the concentrically ridged
but elongate and thin-shelled Corbicula (s.l.) arnoudii, all
the Pourcy corbiculids have more or less smooth shells
and are convergent with the living genera Polymesoda
and Geloina which inhabit estuarine swamps and tropi-
cal/subtropical mangroves and lagoons respectively.
This is taken as an indication that these smooth Ceno-
zoic corbiculids were brackish water species, some of
which might require new genera. Here we assign these
species to Corbicula s.l. The fragile but well-preserved
shells of Corbicula (s.l.) arnoudii were frequent at Pour-
cy although the relationships of this species are not fully
known. It bears some resemblance to the genus Batissa
which today lives in low salinities of the upper estuarine
channel of the Abatan River (Lozouet & Plaziat, 2010).
All corbiculid species at Pourcy were represented by
abundant growth series from the smallest juveniles up-
wards. Shells of the relatively uncommon Tellinocyclas
tellinoides were mostly rather waterworn. This species
appears to have a limited stratigraphic range, occurring
in the Sables de Sinceny and also in the Blackheath For-
mation in England which may be considered more or less
coeval with Pourcy on the basis of molluscs, although
mammal-based correlation is not yet certain.
Corbulidae
The single Varicorbula species, occurring in large num-
bers at Pourcy, approaches the Paleocene-Recent genus
Lentidium in morphology, and is presumed to have had a
similar ecology. This in turn resembles Potamocorbula,
in the Abatan River as noted by Lozouet & Plaziat (2010)
who recorded large colonies of Potamocorbula living su-
percially buried in marginal sandbanks of the estuarine
channel in divergent salinities of between 5 and 34‰.
More strongly ornamented Varicorbula species were char-
acteristic of euhaline environments in the Eocene.
Neritidae
Two extant genera are represented at Pourcy. Neritodryas
today lives on the trunks of waterside trees in the lower
(freshwater) reaches of tropical rivers and streams. The
Clithon species at Pourcy are all referred to the subge-
nus Pictoneritina which typically lives intertidally in
lagoons and estuaries, although other species of Clithon
may extend some distance upstream to freshwater. The
species diversity at Pourcy is comparable with that oc-
curring at Abatan and in other island streams in the Indo-
Pacic region, and the Pourcy species could all have been
contemporaneous. However it is notable that the rare C.
barbei also occurs exclusively with Potamides funatus
in the underlying Argiles à Lignites d’Epernay at Saran
near Epernay while the common Neritoplica uniplicata
and Clithon sobrinum are found in other beds of the same
formation at Mont Bernon, Epernay (Laurain et al., 1983)
and at Mutigny to the east of Pourcy (pers. obs. ST).
Melanopsis
Although species of Melanopsis in Europe today are re-
stricted to freshwater streams and lakes, their morpho-
logically similar relatives in the Indo-west Pacic (New
Caledonia, New Zealand) are often abundant in the brack-
ish tidal estuaries of streams and rivers, sometimes in close
association with mangroves (Franc, 1956; pers. obs. ST).
Tympanotonos/Potamides
Living Tympanotonos occurs only in West Africa where it
inhabits estuarine mudats and mangrove areas both dis-
tally and proximally. We follow Reid et al. (2008, p.692)
who considered that all Cenozoic Tympanotonos species
should be referred to Potamides (type species: Cerithium
lamarckii Brongniart, 1810; Oligocene, France). This has
not been widely accepted, perhaps owing to the fact that
several other genus-level taxa have been proposed within
this group, mostly poorly dened. For this reason we use
Potamides pending a full phylogeny of the group tak-
ing account of their ontogeny (work in progress). This is
also pragmatic as the name Potamides Brongniart, 1810
would replace Tympanotonos Schumacher, 1817 if the two
were considered synonymous. Although Cenozoic Pota-
mididae were clearly not restricted to mangrove settings,
their constant presence in the underlying early Ypresian
laminated clays suggests a similar palaeoenvironment to
1 Bieler et al. (2010) thought the available senior name, Cyrenidae
(Gray, 1840), had been used and so could not be considered a nomen
oblitum, but Article 40.2 of the ICZN Code states ‘If, however, a fam-
ily-group name was replaced before 1961 because of the synonymy
of the type genus, the substitute name is to be maintained if it is in
prevailing usage.’ We regard the family name Corbiculidae Gray, 1847
to have been in prevailing usage throughout most of the 20th century,
and that it would be confusing to revert to a disused family name based
on a junior synonym, so we therefore prefer to conser ve Corbiculidae
(ICZN 1999).
Cainozoic Research, 15(1-2), pp. 155-180, October 2015 175
that of the morphologically analogous living Tympano-
tonos. Other modern potamidids are always associated
with sheltered habitats of reduced salinity, particularly in
areas with mangroves (Reid et al., 2008).
Hydrobiidae
The abundant small, mostly smooth species are here re-
ferred to the Hydrobiidae s.l. pending an in-depth taxo-
nomic assessment. The intraspecic variation is better
judged from populations found at specic horizons in the
underlying clays.
Molluscan feeding guilds
Overall, there is a good resemblance between the feeding
guild compositions of the Eocene Pourcy and the modern
Abatan mollusc faunas, with the caveat that no stand-
ardised collecting has been undertaken (Figs 108, 109).
About half of each fauna is composed (both in species
diversity and in abundance) of herbivores/detritivores.
Carnivores are more common in the Abatan fauna at the
expense of suspension feeders. In both Pourcy and Aba-
tan samples the diversity of carnivores is greater, but they
are not abundant.
The seven samples from the Abatan river taken by Lo-
zouet & Plaziat (2010) spanned a transect from the river
channel to the open bay mouth. Predatory carnivores
were absent in the riverine and upper estuarine samples
(Fig. 109). In species numbers the herbivores dwindled at
the expense of carnivores towards the marine end of the
transect. The abundance data did not show a clear trend.
Figure 108. Mollusc feeding guilds of Pourcy and Abatan.
H: herbivores/detritivores, CP: predatory carnivores, CB:
browsing carnivores, SU: suspension feeders, DS: deposit
feeders, CD: chemosymbiontic feeders.
Figure 109. Feeding guilds of the Pourcy and the entire Abatan
faunas, and those of the seven sample stations from Abatan
that represent riverine to marine settings. Abatan data from
Lozouet & Plaziat (2010). H: herbivores/detritivores, CP:
predatory carnivores, CB: browsing carnivores, SU: sus-
pension feeders, DS: deposit feeders, CD: chemosymbion-
tic feeders. The feeding guild is given for species diversity
(upper graph) and relative abundance (lower graph) data.
The feeding guild composition of the Pourcy fauna
showed the highest resemblance to middle, and lower
estuarine and mouth bay faunas of Abatan. However,
the much better resemblance of the feeding guilds of the
Pourcy fauna with the entire Abatan data set suggests
that Pourcy represents an environmentally (time) aver-
aged fauna. This is also shown by the admixture of appar-
ently ecologically incompatible species at Pourcy, such as
freshwater-oligohaline taxa, (e.g. sphaeriid bivalves and
viviparid and pulmonate snails) with mesohaline-polyha-
line taxa, (e.g. oysters, mussels and turritellid snails).
Laurain et al. (1983) listed the molluscs from the fos-
siliferous sands and clays of beds 16-32 of the Epernay
Formation at Mont Bernon (Marne). These beds occupy
the lower two-thirds of the formation. Signicantly 20
of the 23 species recorded also occur at Pourcy. Tuniot
(1902) gave other occurrences of the species he listed
from Pourcy. Signicantly, none of the species listed
were unequivocally indicators of a Cuisian date except
perhaps Barbatia modioliformis (also at Sinceny and
Cuise-la-Motte and higher strata) which was also one of
the few species usually regarded as indicating euhaline
conditions (although Tuniot, 1902 also recorded it from
‘Les Lignites’). Cyrtopleura orbignyana, occurring at
Sinceny and Cuise-la-Motte, was also identied from a
176 Spijkerman et al. Pourcy: preliminary assessment of an early Eocene NW European tropical coastal environment
few fragments at Pourcy and is regarded as a marine indi-
cator. Eopleurotoma pourcyensis, which would normally
be considered a fully marine species, was also recorded
from the Argiles à Lignites at Mailly-Champagne and
Rilly by Bellevoye (1907) who remarked that it was truly
a Sparnacian species. In the lower beds of the Epernay
Formation examined at Saran (Marne) to the southeast
of Pourcy it was found that Potamides funatus almost ex-
clusively occupied one stratum of the ‘lignites’ while the
anged morph of Potamides turris (Fig. 45) was similarly
abundant at a lower horizon. This adds support to the the-
ory that the Falun de Pourcy with its confusing intraspe-
cic variation among the Potamididae is in fact a mixture
that includes material reworked from underlying levels.
The clays of Mutigny to the east of Pourcy contained an
impoverished but similar mix of subspecic variants to
those at Pourcy. Duprat et al. (2010) placed Mutigny be-
tween the ‘upper and lower levels’ at Pourcy.
The Pourcy fauna has a dissimilar feeding composition
from those of other fossil coastal tropical associations
studied so far. A Miocene Indo-Pacic sea grass fauna
(Reich et al., 2014) shows a much higher abundance of
the herbivore/detritivore guild. Instead, a coral-carpet as-
sociated fauna from the same region shows dominance
of carnivores (A. Kusworo, pers. comm., 2013). A Pleis-
tocene uviolacustrine to high estuarine environment
from Trinil, Java (Joordens et al., 2009) is dominated
(>90% in abundance) by thiarid snails (grazers/detriti-
vores). The accompanying mollusc fauna included Cor-
bicula (s.str.) and indicated the upper reaches of a river or
stream although neritid snails were lacking in the Trinil
fauna. Overall the composition of molluscan feeding guilds
at Pourcy was broadly most similar to that of the modern
Indo-Pacic mangrove-estuarine setting of Abatan.
Vertebrates
The shark teeth were predominantely of juvenile animals,
which indicates a possible mangrove setting (T. Bor, pers.
comm.) The bony sh genera Amia and Lepisosteus are
compatible with a tropical or subtropical, brackish to
freshwater environment with shallow lakes and swamps
(V.W.M. van Hinsbergh, pers. comm., 2013; Wilson, 1982;
Winkler, 1983). The turtles too, indicate a tropical envi-
ronment with estuarine and terrestrial taxa represented
in the material (De Broin, 1977; Buffetaut et al., 1982;
Davenport, 2011; Jimenez-Fuentes et al., 1994). The al-
ligatorid crocodile species implies brackish conditions
and the crocodyliid species implies a freshwater environ-
ment (Davenport, 2011). The presence of the bird genus
Gastornis suggests a warm to subtropical coastal lowland
with alluvial oodplains, well-vegetated swamps and open
savannas (Andors, 1991; Mustoe et al., 2012).
Among the mammals, the relatively large number of pri-
mates is remarkable, with seven genera. This is a strong
indication of a warm, relatively aseasonal climate result-
ing in the availability of seeds and fruits during the entire
year (van den Hoek Ostende, pers. comm.). Genera found
at Pourcy were mainly fruit- and seed-eaters, just like
the eight genera of rodents of this locality. The hoofed
animals were represented by Condylarthra and primi-
tive Artiodactyla and Perissodactyla. These were usu-
ally small forms adapted to live in or among trees. Some
representatives belonged to groups that are now extinct,
such as the Tillotheriidae and the Coryphodont idae. The
tillothere Franchaius is considered to have been a root
and tuber eater (Agusti & Anton, 2002) and the semi-
aquatic Coryphodon had a similar diet. The only carni-
vore in the assemblage, the creodont Palaeo nictis, hunted
in trees. Overall the mammals represent a relatively hu-
mid, forest environment.
This assessment of the palaeoecology of the mammal
fauna raises the question of their relation to an estuarine
mangrove environment as deduced from the other fossils,
and whether they might be derived from earlier terrestrial
deposits. Hooker (1996) originated a series of Paleocene-
Eocene mammal zones dened by rst and last appear-
ances of various taxa. Zone PEI was dated as latest Paleo-
cene dened by the last appearances of various species
including Landenodon woutersi, Paschatherium dolloi,
Microhyus musculus and Phenacodus teilhardi. Zone PEII
included the rst appearance of Neomatronella luciannae
and was dated as early Eocene. These species all occur
together in the Falun de Pourcy which has caused prob-
lems with dating the deposit (see Hand et al., 2015). Cave-
lier (1987) also commented on this phenomenon and cited
earlier similar observations. Mammals of zone PEI would
not be expected to occur in the Epernay Formation, except
perhaps in reworked material at the base, so their presence
at Pourcy in the upper part of the formation is enigmatic.
Other fossils found at Pourcy conrm the general palaeo-
ecology as deduced from molluscs and vertebrates. Four
species of ostracods, Cytheridea sp., Clitrocytheridea
canceratica Apostrolescu, 1956, Vetustocytheridea gui-
trancourtensis Apostrolescu, 1956 and Hemicyprideis
sp. (identications by C.M. Pirkenseer, Fribourg, Swit-
zerland) suggest a brackish coastal, lagoonal or estuarine
environment (C.M. Pirkenseer, pers. comm., 2012). Rare
occurrences of serpulid worms (Ditrupa plana (Sowerby,
1815) and Rotularia bognoriensis (Mantell, 1822) indi-
cate lagoonal to open marine settings (Morton & Harper,
2009; Morton & Salvador, 2009). In England Ditrupa
plana rst appears in the Tilehurst Member of the Har-
wich Formation while Rotularia bognoriensis appears
in the lower part of the London Clay Formation, both in
fully marine facies. The echinoid Scutellina lenticularis
(Lamarck, 1816), is characteristic of marine environ-
ments (J. Eberson, Utrecht, pers. comm. 2008). In addi-
tion a few poorly preserved freshwater charophytes and
a single small example of the foraminifer Nummulites
planulatus were found. This species reached its acme
in the mid-late Ypresian (Cuisian) which is usually re-
garded as its rst appearance, although it is also recorded
from Sinceny (Aisne) (MNHN-GG-GG2004-77213 Coll.
Abrard, Soyer; MNHN-GG-GG2004-77221 Collection
Bonnet) which may be tentatively correlated with the Fa-
lun de Pourcy on the basis of molluscs, although mammal
records are equivocal (Hooker, 2015).
Cainozoic Research, 15(1-2), pp. 155-180, October 2015 177
In conclusion, the invertebrate faunas from Pourcy rep-
resent early Eocene tropical coastal settings probably
with mangroves. They contain indicators for full ma-
rine, euryhaline and freshwater as well as terrestrial set-
tings. The forest environment, which may have been the
source of the mammals, suggests a degree of transport
and certainly some reworking. The same is probably true
of the molluscs as the intraspecic variation seen in the
common brackish water groups, mentioned above, is far
greater than would be expected among living populations
in any one locality today.
A landscape reconstruction made for the North Sea Basin
Eocene provided in Fig. 110 shows a palaeoenvironment
probably similar to that of Pourcy.
Figure 110. Reconstructed mangrove landscape in the Ypresian of the southern North Sea Basin that at the time included also the
Paris Basin. Drawing by E.J. Bosch (Naturalis Biodiversity Center, Leiden).
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180 Spijkerman et al. Pourcy: preliminary assessment of an early Eocene NW European tropical coastal environment
Class Bivalvia
Family Arcidae 13 SU M
Family Noetiidae 96 SU M
Family Mytilidae 139 SU M/B
Family Ostreidae 268 SU M
Family Anomiidae 87 SU M
Family Lucinidae 42 CD M
Family Psammobiidae 2 D M
Family Corbiculidae 1943 SU B/(F)
Family Pisidiidae 1 SU F
Family Veneridae 19 SU M
Family Myidae 24 SU M
Family Corbulidae 855 SU M/B
Family Pholadidae 13 SU M/B/(F)
Class Gastropoda
Family Neritidae 844 H B/F/(M)
Family Viviparidae 2 H F
Family Ampullinidae 110 H M
Family Potamididae 797 H B/M
Family Batillariidae 32 H B/M
Family Pachychilidae 460 H B/F
Family Melanopsidae 834 H B/F
Family Thiaridae 11 H B/F
Family Turritellidae 49 SU M
Family Hydrobiidae (s.l.) 572 H B/F
Family Naticidae 113 CP M
Family Epitoniidae 2 CP M
Family Turbinellidae 11 CP M
Family Buccinidae 53 CP M
Family Pseudolividae 21 CP M
Family Turridae 10 CP M
Family Cornirostridae 163 H M
Family Pyramidellidae 85 P M/(B)
Family Ringiculidae 3 CP M
Family Cylichnidae 2 CP M
Family Carychiidae 2 CP T/(F)
Family Physidae 2 H F
Family Planorbidae 33 H F
Appendix 1
Information about abundance (number of specimens in the
studied collections), feeding-behaviour and salinity prefer-
ences of the mollusc families. Only material from the studied
collections is included (not taxa recorded in literature only or
additional material in private collections). Salinity preferences
are estimated and only an approximation.
Abbreviations for feeding ecology:
SU = suspension feeder; D = deposit feeder; CD = chemosym-
biontic deposit feeder; H = herbivore/detritivores; P = parasite;
CP = predatory carnivore. Salinity: F = freshwater; B = brack-
ish (stenohaline/euryhaline); M = marine (euhaline); T = ter-
restrial.
Appendix 2
Additional records of sh remains from the two sites at Pourcy
in the Allan Lawson collection (Mill Hill, London, England),
which are not included in the statistics.
Abundance Feeding Salinity
(n) behaviour preference
Synechodontidae
Synechodus sp. 1 tooth (rolled)
Heterodontidae
Heterodontus sp. 1 tooth (rolled)
Carcharhinidae
Physogaleus secundus (Winkler, 1874) 6 teeth
Odontaspididae
Striatolamia macrota (Agassiz, 1843) 1 tooth
Jaekelotodontidae
Palaeohypotodus rutoti (Winkler, 1874) 1 basal fragment
Scyliorhinidae
Scyliorhinus gilberti (Casier, 1946) 1 tooth
Scyliorhinus sp. 1 tooth
Rhinobatidae
Rhinobatos bruxelliensis (Jaekel, 1894) 1 tooth
Dasyatidae
Dasyatis jaekeli Leriche, 1905 7 teeth
Jaquhermania duponti (Winkler, 1874) 1 tooth
Hypolophidae
Hypolophodon sylvestris (White, 1931) 1 tooth (crown)
Lepisosteidae
Lepisosteus suessionensis Gervais, 1852 34 teeth, 20 scales
Amiidae
Amia sp. 1 tooth
Albulidae
Pterothrissus tardinensis (Leriche, 1908) 17 otoliths
Synodontidae
Argentina abbatiae Stinton, 1965 4 otoliths
Serranidae
Polyperca serranoides Stinton, 1965 10 otoliths
serranid sp. 1 2 otoliths
PERCOIDEI
Anthracoperca cf. siebergi Voigt, 1934 4 otoliths
LABROIDEI
Diaphyodus sauvagei (Leriche, 1900) 6 palate fragments
