Juraphyllites mimatensis (D'ORBIGNY, 1845 ... - Palaeodiversity

Palaeodiversity 1: 133–140; Stuttgart, 30.12.2008. 133
Juraphyllites mimatensis (d’Or b i g n y, 1845) (Ammonoidea:
Phylloceratida), a Tethyan immigrant in the Upper Pliensbachian of
Franconia (Jurassic, Southern Germany)
He l m u t Ke u pp & gü n t e r Sc H w e i g e r t
Abstract
Two well-preserved small-sized specimens of phylloceratoid ammonites are reported from the Late Pliensbachian
Spinatum Zone of Franconia. They are assigned to Juraphyllites (cf.) mimatensis (d’Orb i g n y, 1845) and regarded
as Tethyan immigrants. Due to their very similar early ontogenetic stages it seems impossible to distinguish
tiny juveniles of Juraphyllites mimatensis (d’Orb i g n y, 1845) from those of contemporaneous Meneghiniceras eximium
(v. Ha u e r).
Key words: Phylloceratina, Early Jurassic, palaeobiogeography, immigration, Franconia, Germany.
Zusammenfassung
Zwei gut erhaltene kleinwüchsige Exemplare phylloceratoider Ammoniten werden aus der Spinatum-Zone des
Ober-Pliensbachium von Franken beschrieben. Sie werden zu Juraphyllites (cf.) mimatensis (d’Orb i g n y, 1845) gestellt
und als tethyale Einwanderer angesehen. Wegen ihrer sehr ähnlichen frühontogenetischen Stadien erscheint
es praktisch unmöglich, sehr kleine Individuen von Juraphyllites mimatensis (d’Orb i g n y, 1845) von solchen des
gleich zeitig vorkommenden Meneghiniceras eximium (v. Ha u e r) zu unterscheiden.
1. Introduction
The dark claystones of the Upper Pliensbachian
Amaltheenton Formation of Southern Germany, particularly
of Franconia, yield a relatively low diverse ammonite
fauna, generally dominated by the Subboreal family of the
Amaltheidae. Rare immigrations of various Tethyan or
Euroboreal ammonite taxa seem to be restricted to short
phases of sea level highstands, as recently pointed out by
Sc H w e i g e r t (2005) and Ke u pp & Sc H O b e r t (2008). These
exotic ammonites attracted special attention and were noted
by numerous former authors (see Sc H w e i g e r t 2005 for
references; ma i S c H 2007).
Soon after the publication of some findings of juvenile
phylloceratids from the clay-pit of Buttenheim in Franconia
(Ke u pp & Sc H O b e r t 2008), two somewhat larger specimens
were found by the amateur collector JO H a n n Sc H O ­
b e r t (Hirschaid). Both specimens exhibit diameters of 30
and 43 mm, respectively, and are assigned to the unkeeled
juraphyllitid taxon Juraphyllites (cf.) mimatensis
(d’Orb i g n y, 1845). This taxon, which is of Tethyan origin,
was previously unknown from the Pliensbachian of Southern
Germany. The new findings, which question earlier
determinations of juveniles, stimulated this study.
Acknowledgements
JO H a n n Sc H O b e r t (Hirschaid) is thanked for his generous
donation of his interesting findings for this study. The journals’
referees JO a c H i m bl a u (Gießen) and cH r i S t i a n me i St e r (Genève)
are thanked for their advice and suggestions.
2. Provenance of the studied material
The Middle Liassic Amaltheenton Formation (Upper
Pliensbachian) of Franconia crops out in the western foreland
of the Northern and Middle Franconian Alb and is
exposed in several clay-pits. Today most of them are abandoned
(e. g., Kalchreuth: ri c H t e r 1993; Marloffstein:
plü c K e bau m 1985; Sc H m i d t­Ka l e r 2004; Reichenschwand:
Haa r l ä n d e r 1961; Schnaittach: vO i g t 1968; Unterstürmig:
ri c H t e r 1977, 2000; plü c K e bau m 1985). The only
still active clay-pit of the Liapor Company in this area lies
in the Oberfranken district, about 1 km SSE of Buttenheim.
This clay-pit substitutes the former clay-pit of Unterstürmig
which was located only 1.5 km to the south.
The latter outcrop became popular by the occurrence of
well-preserved white-coloured amaltheids bearing their
original aragonitic shells (du l l O 1981). The stratigraphic
sections exposed in both clay-pits are almost identical
(ri c H t e r 2003; HOf f m a n n et al. 2007). In the former claypit
of Unterstürmig especially the upper part of the Spinatum
Zone of the Upper Pliensbachian and the Lower Toarcian
was exposed; only from time to time expanding
downwards to the Apyrenum Subzone of the Upper Pliensbachian.
In contrast, the clay-pit near Buttenheim exposes
primarily beds of the Gibbosus and Apyrenum subzones,
expanding upwards during the progress of the last two
years to the upper part of the Spinatum Zone and the base
of the Lower Toarcian Posidonia Shale (Fig. 1). Also in the
clay-pit of Buttenheim well-preserved ammonites occur

134 pa l a e O d i v e r S i t y 1, 2008
Fig. 1. Geographical position und section of the clay pit near Buttenheim. The horizon where the Juraphyllites mimatensis (d’Orb i g n y,
1845) illustrated in Fig. 3 was found is marked by an asterisk.

with their primary aragonitic shells, sometimes, especially
within beds of the Apyrenum Subzone, showing a colour
pattern formed by dark, sub-concentric dotted spiral lirae
(ri c H t e r 2003; Ke u pp 2005; Ke u pp & Sc H O b e r t 2008).
Within a collection of ca. 17.000 ammonites from the
Upper Pliensbachian of Buttenheim there are only less than
ten phylloceratids, most of them juveniles. The first phylloceratid
specimen described in this study was originally
included in a small calcareous concretion. It was collected
by J. Sc H O b e r t (Hirschaid) in early March 2008 in the
higher section of the clay-pit of Buttenheim. It came from
undisturbed sediments 20 cm below the main fossiliferous
horizon of the upper Spinatum Zone (= Hawskerense Subzone).
The predominant ammonites of this level are the
different morphotypes of Pleuroceras spinatum (Br ug
u i è r e) which occur mainly in the directly overlying beds.
The second specimen was found loose also by J. Sc H O ­
b e r t, together with dispersedly distributed amaltheids of
the Apyrenum Subzone in early spring 2007.
The studied ammonites are housed in the Institute of
Geological Sciences, Freie Universität Berlin, collection
Ke u pp, numbers MAn-3111 and MAn-3115.
3. Systematic palaeontology
Suborder Phylloceratina arK e l l, 1950
Superfamily Phylloceratoidea Zi t t e l, 1884
Family Juraphyllitidae arK e l l, 1950
Genus Juraphyllites mü l l e r, 1939
Ty pe species: Phylloceras diopsis ge m m e l l a rO, 1884.
K e u p p & S c H w e i g e rt, j u r a p h y l l i t e s m i m a t e n s i s in S O u t H e r n g e r m a n y 135
Remarks. – In several recent publications dealing
with Juraphyllitidae, the genera Meneghiniceras Hya t t,
1900, Harpophylloceras Spa t H , 1927, and Juraphyllites
mü l l e r, 1939 were considered to be synonymous or of
subgeneric rank (e. g., me i St e r 1989; ma c c H i O n i 2001;
ma c c H i O n i in pav i a & cr e S ta 2002; ma c c H i O n i & me i St e r
2003). Indeed, Harpophylloceras can be easily included in
Meneghiniceras (Sc H w e i g e r t 2005), but the latter differs
from other Juraphyllitidae by the presence of a keel in the
adult stage. Therefore we maintain the genus Juraphyllites
for the forms lacking a keel.
Juraphyllites mimatensis (d’Orb i g n y, 1845)
Figs. 2–3, ? 4
* 1845 Ammonites Mimatensis, d’Orb i g n y, 1844. –
d’Orb i g n y, p. 344, pl. 110, figs. 4–6.
1856 Ammonites Mimatensis, d’Orb., 1845. – Oppe l,
p. 252.
1929 Rhacophyllites spec. ex aff. mimatensis Or b. –
gu g e n b e r g e r , p. 255, pl. 10, fig. 5, pl. 11, fig. 6,
pl.12, fig. 10.
? 1930 Rhacophyllites cf. mimatensis (d’Orb i g n y). –
mO n e S t i e r , p. 8, pl. 11, fig. 1.
1934 Rhacophyllites mimatensis (d’Orb i g n y). – mO n e St
i e r, p. 16, pl. 5, figs. 17, 19, 24–30, 34–35.
1934 Rhacophyllites libertus (ge m m e l l a rO). – mO n e St
i e r, p. 17, pl. 6, figs. 1, 6.
? 1934 Rhacophyllites mimatensis (d’Orb i g n y) var. vinensis.
– mO n e S t i e r , p. 17, pl. 5, figs. 36–37.
pars 1963 Rhacophyllites mimatensis (d’Orb i g n y). – Sg u a Z­
Z O n i, p. 46, pl. 21, fig. 10, non fig. 4.
? 1970 Juraphyllites libertus (ge m m e l l a rO). – br a m b i ll
a, p. 262, pl. 55, fig. 2.
1977 Juraphyllites mimatensis (d’Orb i g n y). – wi e d e nm
a y e r , p. 36, pl. 1, figs. 3, 8b.
Fig. 2. Juraphyllites mimatensis (d’Orb i g n y, 1845), holotype, lost; reproduction of d’Orb i g n y’s illustration; Upper Pliensbachian; Mt.
Mimat, Dept. Lozère, France. Diameter 42 mm.

136 pa l a e O d i v e r S i t y 1, 2008
Fig. 3. Juraphyllites mimatensis (d’Orb i g n y, 1845); Pliensbachian, upper part of the Spinatum Zone, Hawskerense Subzone; Buttenheim,
northern Bavaria, Germany; MAn-3111. 29.2 mm in diameter. – A. Lateral view. B. Ventral view. C. Oblique view showing
the large missing shell fragment due to a presumed crustacean attack. D. Close-up of Fig. 3A showing the suture line.
1989 Juraphyllites mimatensis (d’Orb i g n y). – me i St e r,
p. 30, pl. 2, fig. 10, pl. 3, figs. 4, 17a, b.
1994 Juraphyllites mimatensis (d’Orb i g n y). – JO ly in
fi S c H e r, p. 98, fig. 31, pl. 27, figs. 3a–c, 4.
2000 Juraphyllites mimatensis (d’Orb i g n y, 1845). –
JO ly, p. 29–30, figs. 41–43, pl. 3, fig. 6a, b.
Types: Juraphyllites mimatensis was first described by
d‘Orb i g n y (1865) from the Upper Pliensbachian of Southern
France and named after the type locality, the Mont Mimat south
of Mende (Dept. Lozère). Because of the apparent lost of the
type specimen originally housed in the re n a u x collection in
Montpellier, JO ly (in fi S c H e r 1994) designated a neotype, a
mould coming from the middle Upper Pliensbachian (Margaritatus
Zone) of Mt. Guilhaumard near Tournadous. This neotype,
however, is much less complete than the lost holotype, following
the illustration by d’Orb i g n y (1845) (Fig. 2).
Occur rence: From different localities of the Grands Caus-

ses (Southern France, e. g., d’Orb i g n y 1845; Oppe l 1856; mO n e St
i e r 1934; me i St e r 1989; JO ly in fi S c H e r 1994, JO ly 2000),
Turkey (gu g e n b e r g e r 1929); ? Morocco (mO n e S t i e r 1930), ?
West-alpine Italy (br a m b i l l a 1970), South-alpine Switzerland
(wie d e n m a y e r 1977), Apennines (Sg u a Z Z O n i 1963), and Franconia
(herein).
Stratigraphic range: Late Pliensbachian (Stokesi to
Hawskerense subzones), possibly already present in the upper
Early Pliensbachian (Davoei Subzone), see wi e de n m ay e r (1977).
In the Causses region this taxon is only reported from the Subnodosus
to Gibbosus subzones (me i St e r 1989), but unknown yet
from the Spinatum Zone.
Descriptions. – The first and smaller one of the two
studied specimens (MAn-3111) measures 29.2 mm in diameter.
The phragmocone of this completely preserved
specimen is slightly compressed by compaction processes,
whereas its body chamber (last ½ whorl) does not show
any diagenetic deformation (Fig. 3). However, on one of
the flanks, the body chamber has lost a large angular piece
of the shell affecting the entire flank and reaching from
the peristome about 16 mm backwards. Obviously this
specimen was attacked by a predator, possibly a benthic
crustacean. Due to this large artificial opening of the body
chamber, the measurements characterizing the shell morphology
was taken along the smaller diameter of 24.2 mm
(see below). On the surface of the mould some relics of the
white aragonitic shell material are still preserved. The
slightly compressed phragmocone shows the characteristic
juraphyllitid suture line (Fig. 3D) which matches perfectly
with that of the lost holotype illustrated by d’Orb i g n y
(1865; Fig. 2) and the illustrations of the suture lines of the
neotype and additional material from the Causses provided
by JO ly (in fi S c H e r 1994) and JO ly (2000). Moreover,
also the measurements fall within the size ranges of the
specimens from the Causses (JO ly 2000; see Tab. 1). The
umbilicus of the specimen from Buttenheim is only slightly
smaller, and the cross-section of the whorl is insignificantly
broader. Surprisingly, the holotype (Fig. 2) is the
closest one to our present finding although later authors
stated that the illustration by d’Orb i g n y could be somewhat
generalized.
K e u p p & S c H w e i g e rt, j u r a p h y l l i t e s m i m a t e n s i s in S O u t H e r n g e r m a n y 137
Tab. 1. Measurements of specimens related to Juraphyllites mimatensis (d = diameter; u = umbilicus; wh = whorl height; ww = whorl
width).
MAn-3111
(Fig. 3)
MAn-3115
(Fig. 4)
specimen
Neotype
(JO ly in fi S c H e r 1994)
Coll. JO ly no. 99-1 Cornus
(JO ly 2000)
Coll. JO ly no. 140-1 Puech
(JO ly 2000)
d u u/d wh wh/d ww ww/d ww/wh
(mm)
29.2
(mm) (%) (mm) (%) (mm) (%) (%)
24.2 6.6 27 11.6 48 7.9 32.5 68
43 6.8 15.8 24.0 56 12.1 28 50
34.5 10 29 16 46 9.8 28 61
27.3 8.8 32 12.3 45 7.3 27 59
22 6.8 31 9.7 44 6 27 62
On the body chamber simple ribs occur, running
slightly convex over the flattened flanks, and disappear on
the rounded venter. Immediately behind the peristome, the
ribs tend to become fastigate. The almost smooth venter
does not show any indication of a keel. Within the umbilicus
about six constrictions per whorl are discernible which
seem to disappear successively after a diameter of ca.
18 mm.
The incompletely preserved second specimen (MAn-
3115, Fig. 4) has a diameter of ca. 43 mm. It exhibits a
nearly completely preserved and uncrushed body chamber
(last half whorl). The phragmocone has been collapsed by
compaction processes and was lost along the first half of
the outer whorl, but is still visible within the open umbilicus.
The weak ribs run more or less radially over the flattened
flanks of the body chamber and cross the rounded
venter along the last ¼ whorl in a fastigate style. Within
the open umbilicus, up to six constrictions per whorl are
discernible. The body chamber itself does not show any
constrictions. The morphology differs from the previously
described specimens of Juraphyllites mimatensis especially
in the measurements of the umbilicus, which is significantly
narrower, with a relative width of about 16 %
versus 27–32 %, and the cross-section of the whorl which
appears to be higher (ww/wh = 50 % versus 59–68 %).
Only the umbilical elements of the last suture line are preserved
just at the beginning of the body chamber showing
a characteristic juraphyllitic pattern formed by short amplitudes.
From the suture line aspect, this specimen is
clearly distinguishable from the contemporaneous phylloceratid
genus Zetoceras. In the latter, a similarly narrow
umbilicus is developed, but its suture line is much more
complex, with high amplitudes (see JO ly 2000). The suture
line, ribbing sculpture, and the lacking of a keel of this
specimen correspond with Juraphyllites mimatensis. However,
due to the significantly smaller umbilicus and the
higher whorl-section we prefer a cf.-determination of this
specimen.

138 pa l a e O d i v e r S i t y 1, 2008
Fig. 4. Juraphyllites cf. mimatensis (d’Orb i g n y, 1845); Pliensbachian, Spinatum Zone, ? Apyrenum Subzone; Buttenheim, northern
Bavaria, Germany; MAn-3115. – A. Lateral view. B. Ventral view. – Scale: 2 cm.
4. Palaebiogeography of Juraphyllites mimatensis
(d’Or b i g n y, 1845)
Without any doubt Juraphyllites mimatensis represents
a Tethyan faunal element of the Late Pliensbachian which
immigrated only during sea-level highstands and favourable
currents into adjacent epicontinental basins, such as
the Causses area of Southern France and reached even
Southern Germany (see above). The records of this species
in true Tethyan deposits are still remarkably scarce and
seem to be restricted to the Western Tethys. However, it is
not unlikely that this species is much more widespread but
still unrecognized (see wi e de n m ay e r 1977: 37), because in
early ontogenetic stages this form is hardly distinguishable
form other juraphyllitids such as Meneghiniceras eximium
(v. Ha u e r) and Juraphyllites libertus (ge m m e l l ar
O).
5. Previously described juvenile Phylloceratina from
the Upper Pliensbachian of Southern Germany
Qu e n S t e d t (1885) described and illustrated tiny phylloceratids
from the Amaltheenton Formation (“Lias δ”) of
Gross-Eislingen (today: Eislingen) in Swabia as “Ammonites
tortisulcoides”. Other records assigned to this species
have been recorded from Southern Germany by Zw i e­
Se l e (1898), en g e l (1908), ur l i c H S (1977), plü c K e bau m
(1985), and Sc H l e g e l m i l c H (1976). On the other hand,
HO w a r t H (1976) reported on a single excellently preserved
finding of the keeled juraphyllitid Meneginiceras eximium
(Ha u e r , 1854) var. lariense (me n e g H i n i, 1874) from the
Lower Toarcian of the British Yorkshire coast. The innermost
whorls of Meneghiniceras have not developed the
characteristic ventral keel of later ontogenetic stages and
hence they correspond morphologically well with the
small and partly still unkeeled specimens of Ammonites
tortisulcoides Qu e n S t e d t. As a consequence, Sc H w e i g e r t
(2005) interpreted the taxon Ammonites tortisulcoides
Qu e n S t e d t, 1885 as a subjective junior synonym of Meneghiniceras
eximium (v. Ha u e r , 1854) due to the tendency
to form a ventral keel with increasing diameters in most
of his specimens of Ammonites tortisulcoides under study.
Interestingly, mO n e S t i e r (1934) interpreted a specimen
from Swabia figured by pOm pe c K J (1893, pl. 1, figs. 10–11)
as belonging to d’Orb i g n y’s species mimatensis, but he
overlooked the keeled venter of this specimen. The presence
of a keel also in the lectotype of Ammonites tortisulcoides
confirms the supposed synonymy of the two taxa.
Ke u pp & Sc H O b e r t (2008) recorded for the first time several
very small, juvenile phylloceratid phragmocones, all
of them less than 1 cm in diameter, from the Apyrenum
Subzone of Buttenheim. Following the argumentation
given by Sc H w e i g e r t (2005), these innermost whorls were
also determined as juveniles of the juraphyllitid Meneghiniceras
eximium (Ha u e r) although a keel is not developed
in any of these tiny specimens. In Swabia, Meneghiniceras
eximium (v. Ha u e r) occurs as an iteratively immigrating
Tethyan guest in the Late Pliensbachian, but it was
never recorded from the Spinatum Zone.

However, with the herein described specimens of Juraphyllites
mimatensis, a second juraphyllitid species is
now reported which immigrated from the Tethyan into the
epicontinental shallow sea of Southern Germany during
the Late Pliensbachian. The juvenile stages of both taxa
appear to be indistinguishable. The same must be said
from the juveniles of another closely related juraphyllitid
ammonite from the Tethyan Realm, Juraphyllites libertus
(ge m m e l l a r O) (see list of synonyms in fa n t i n i SeS t i n i
1974), which is rather evolute throughout ontogeny and
exhibits coarse ventrolateral ribs in the subadult and adult
stages.
6. Conclusions
In the Late Pliensbachian Tethyan juraphyllitids immigrated
in the Apyrenum and Hawskerense subzones of
Southern Germany and are recorded from Buttenheim in
Franconia. These extremely weak immigration events
have not been recorded yet from Swabia, possibly due to
collecting biases or more marly lithologies which are unfavourable
for ammonite preservation. Besides the juraphyllitids,
represented by Juraphyllites (and cf.) mimatensis
and several undeterminable juveniles, no other Tethyan
immigrants are reported within the ammonite faunas of
the Franconian Upper Pliensbachian absolutely predominated
by amaltheids (Pleuroceras ssp.). An unambiguous
determination of juraphyllitids with diameters of less than
1 cm seems impossible.
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Addresses of the authors:
Prof. Dr. He l m u t Ke u pp, Freie Universität Berlin, Department of Earth Sciences, Palaeontology, Malteserstraße 74-100, House D,
12249 Berlin, Germany
E-mail: keupp@zedat.fu-berlin.de
Dr. gü n t e r Sc H w e i g e r t, Staatliches Museum für Naturkunde, Rosenstein 1, 70191 Stuttgart, Germany
E-mail: schweigert.smns@naturkundemuseum-bw.de
Manuscript received: 26.5.2008, accepted: 29.10.2008.