Content uploaded by Ursula Menkveld-Gfeller
Author content
All content in this area was uploaded by Ursula Menkveld-Gfeller on Sep 27, 2022
Content may be subject to copyright.
Ferrante etal. Swiss Journal of Palaeontology (2022) 141:15
https://doi.org/10.1186/s13358-022-00257-z
RESEARCH ARTICLE
The rst Jurassic coelacanth
fromSwitzerland
Christophe Ferrante1,3*, Ursula Menkveld‑Gfeller2 and Lionel Cavin3
Abstract
Coelacanths form a clade of sarcopterygian fish represented today by a single genus, Latimeria. The fossil record of the
group, which dates back to the Early Devonian, is sparse. In Switzerland, only Triassic sites in the east and southeast of
the country have yielded fossils of coelacanths. Here, we describe and study the very first coelacanth of the Jurassic
period (Toarcian stage) from Switzerland. The unique specimen, represented by a sub‑complete individual, possesses
morphological characteristics allowing assignment to the genus Libys (e.g., sensory canals opening through a large
groove crossed by pillars), a marine coelacanth previously known only in the Late Jurassic of Germany. Morphological
characters are different enough from the type species, Libys polypterus, to erect a new species of Libys named Libys
callolepis sp. nov. The presence of Libys callolepis sp. nov. in Lower Jurassic beds extends the stratigraphic range of the
genus Libys by about 34 million years, but without increasing considerably its geographic distribution. Belonging to
the modern family Latimeriidae, the occurrence of Libys callolepis sp. nov. heralds a long period, up to the present day,
of coelacanth genera with very long stratigraphic range and reduced morphological disparity, which have earned
them the nickname of ‘living fossils’.
Keywords: Sarcopterygii, Actinistia, Libys, New species, Mesozoic, Toarcian, Morphology
© The Author(s) 2022. Open Access This article is licensed under a Creative Commons Attribution 4.0 International License, which
permits use, sharing, adaptation, distribution and reproduction in any medium or format, as long as you give appropriate credit to the
original author(s) and the source, provide a link to the Creative Commons licence, and indicate if changes were made. The images or
other third party material in this article are included in the article’s Creative Commons licence, unless indicated otherwise in a credit line
to the material. If material is not included in the article’s Creative Commons licence and your intended use is not permitted by statutory
regulation or exceeds the permitted use, you will need to obtain permission directly from the copyright holder. To view a copy of this
licence, visit http:// creat iveco mmons. org/ licen ses/ by/4. 0/.
Introduction
Coelacanths, or Actinistia, is one of the two clades of
living sarcopterygian fishes, the second one being the
lungfishes, or Dipnoi. Only known by two extant species,
coelacanths were more diverse in the past since their split
from other sarcopterygians about 420 million years ago,
although they were always very minority compared to
other fish clades.
In the Lower Jurassic, six species of coelacanths are
currently known from marine and freshwater envi-
ronments of Europe, North America, South Amer-
ica and India (Fig. 1A). Most species were recovered
from different localities of Laurasia. e oldest known
Jurassic coelacanth is Diplurus longicaudatus of the
Hettangian-Sinemurian, a Laurasian species of which
the genus is originated in the Carnian (Upper Trias-
sic) and recovered from various freshwater sediments
of United States (Fig.1A1) (e.g., Forey, 1998; Schaeffer,
1948). Laurasian coelacanths are better known in marine
environments of European localities. ese coelacanths
are represented by Holophagus gulo from the Sinemu-
rian of England (Fig.1A2), and the giant Trachymetopon
from the Sinemurian and the Toarcian of Germany, the
Callovian of France and the Kimmeridgian of England
(Fig.1A3) (Cavin etal., 2021a; Dutel etal., 2015; Forey,
1998). Another coelacanth of a poorly defined species
of uncertain affinities, Undina (?) barroviensis has been
described in the Lower Jurassic of England (Fig. 1A4)
(Woodward, 1890, 1891; Forey, 1998). Gondwanian
coelacanths are rarer during the Lower Jurassic and are
represented only by two known species. e marine Ata-
camaia solitaria is the only known coelacanth that lived
on the Paleopacific side of Gondwana (Fig.1A6) during
the Middle to Upper Sinemurian (Arratia & Schultze,
Open Access
Swiss Journal of Palaeontology
Editorial Handling: Daniel Marty
*Correspondence: christophe.ferrante@ville‑ge.ch
1 Department of Earth Sciences, University of Geneva, Rue des Maraîchers 13,
1205 Geneva, Switzerland
Full list of author information is available at the end of the article
15 Page 2 of 20
C.Ferrante et al.
2015). Indocoelacanthus robustus, recovered from the
Kota Formation in India (Fig.1A7), is the only known
coelacanth that lived in the freshwater environments of
Gondwana during the Toarcian (Jain, 1974). is low
taxic diversity of coelacanths during the Lower Jurassic
makes a new occurrence of coelacanth particularly inter-
esting for understanding the evolutionary history of the
group.
In Switzerland, coelacanths are well represented in
the Middle Triassic with Ticinepomis peyeri (Rieppel,
1980), Heptanema paradoxum (Renesto & Stockar, 2018;
Renesto et al., 2021) and a new taxon currently under
description (Ferrante etal., 2017; Rieppel, 1985;Ferrante
etal., work in progress), all recovered from the marine
environment of Monte San Giorgio and Ducanfurgga
(Canton of Ticino and Graubünden, respectively). We
report here the presence of a specimen of a coelacanth
(NMBE 5034072 and 5034073) from the Toarcian, Early
Jurassic, of the Swiss Prealps of the Canton of Fribourg.
is specimen is then the very first coelacanth in Swit-
zerland from another period than the Triassic. Moreo-
ver, this specimen collected in 1870 at the locality of
Les Pueys (Fig. 1C) near Teysachaux (Canton of Fri-
bourg, Switzerland), represents probably the very first
coelacanth fossil found in Switzerland. e identity of
the collector is not assured. It was probably Joseph Car-
dinaux, from Châtel-Saint-Denis (Canton of Fribourg,
Switzerland), a fossil collector and dealer. Joseph Cardi-
naux found and sold many other fossils, notably the first
ichthyosaur from Switzerland (Mennecart & Havran,
2013), also from Teysachaux, purchased by Carl von
Fischer-Ooster, a paleontologist at the Naturhistorisches
Museum Bern (Canton of Bern, Switzerland). When
Carl von Fischer-Ooster incorporated the coelacanth
Fig. 1 Maps showing the localization of the fossiliferous outcrops. A Paleogeographical map showing the continent configuration during the
Toarcian (modified from Scotese, 2014, Map 39) with the distribution of (1) Diplurus longicaudatus, (2) Holophagus gulo, (3) Trachymetopon, (4)
Undina (?) barroviensis, (5) Libys callolepis sp. nov. (6) Atacamaia solitaria and (7) Indocoelacanthus robustus. B Paleogeographic map of the NW Tethys
Ocean (modified from Fantasia et al., 2019), enlargement of B (red square). C Map of Switzerland with the locality of Les Pueys near the Teysachaux
summit (Canton of Fribourg, Switzerland)
Page 3 of 20 15 The rst Jurassic coelacanth from Switzerland
specimen in the collections in 1873–1874, he gave it the
name ‘Macropoma heeri’, but never properly describing
it, probably because he passed away in 1875. is name
is then a nomen nudum according to the International
Code of Zoological Nomenclature. Since its discovery,
the specimen has only been mentioned as ‘Macropoma’
by Hug (1898) in its description of the fauna of Les Pueys
and by Marchant and Pichon (2013) in their general
guide summarizing the essentials of paleontological dis-
coveries in Switzerland.
Geological andpaleogeographic settings
oftheTeysachaux sites
e coelacanth specimen described here was recovered
from a small locality named Les Pueys near the village
of Châtel-Saint-Denis (Canton Fribourg, Switzerland)
(Fig.1C). is locality is situated on the western slope
of the Teysachaux summit, which is on the same moun-
tain ridge than the emblematic Swiss Moléson summit.
e Teysachaux region is best known for another fossil-
iferous site named Creux de l’Ours (Fig.1C) which was
exploited since the nineteenth century. e site of the
Creux de l’Ours has yielded many marine fossils, includ-
ing bivalves, gastropods, ammonites, belemnites, deca-
pods, echinids, trace fossils, plant remains and some rare
vertebrates such as fishes and one almost complete skel-
eton and some remains of ichthyosaurs (Fischer-Ooster
& Ooster, 1870; Furrer, 1960; Huene, 1939; Menkveld-
Gfeller, 1998; Mennecart & Havran, 2013; Weidmann,
1981). In the sites of Creux de l’Ours and Les Pueys, first
fossils were discovered and collected mainly by the ama-
teur collector Joseph Cardinaux, who sold many of his
finds to the Naturhistorisches Museum Bern (Canton of
Bern, Switzerland). From the Creux de l’Ourssite, more
than 400 fossils and from the site of Les Pueys about
a hundred fossils were sold to the Naturhistorisches
Museum Bern. Fischer-Ooster and Ooster (1870), Hug
(1898), Huene (1939) and von der Weid (1960) described
the fauna of these two sites in detail. e name of the
collector who found the coelacanth of Teysachaux is
unknown, but according to the dates of collect (before
1870) and of arriving in collection of the Naturhis-
torisches Museum Bern (1873–74), it is likely that it was
Joseph Cardinaux.
e exact location from which the coelacanth mate-
rial was recovered is not clear but Hug (1898) mentioned
that the fossil beds of Les Pueys are found in a small river.
Prospecting near the place called Les Pueys carried out
by one of us (CF) permitted to find a small valley crossed
by a small river with some fossiliferous layers that could
correspond to the site mentioned by Hug (1898). Com-
parison of the lithology and the invertebrate fossils on
the slab containing the coelacanth specimen with the
material from the Creux de l’Ours bonebeds suggests
that the coelacanth material of Les Pueys is part of the
same lithostratigraphy than the Creux de l’Ours site.
Indeed, on the reverse side of the two slabs with the coe-
lacanth (NMBE 5034072 and 5034073) are various small
coprolites, fish scales and fine fragments of bones, shell
fragments and some poorly preserved evolute forms of
ammonites (Fig.2A, B). e latter are referred to Dac-
tylioceras? (Fig.2A) and Harpoceras renevieri Hug 1898
(Fig.2B) of the families Dactylioceratidae and Hildocer-
atidae, respectively. A bivalve was identified as Goniomya
rhombifera Goldfuss 1840 (Fig. 2C). ose fossils also
resemble to the findings from the same strata of Creux
de l’Ours in terms of the manner in which the shell is pre-
served. e same invertebrate genera and species are also
known from the Creux de l’Ours site (Fig.2D–F).
e Creux de l’Ours section is part of the Stadelgra-
ben Formation from the Nappe of the Préalpes Médianes
Plastiques (e.g., Fantasia et al., 2018; Weidmann, 1993).
Septfontaine (1983) introduced the term Staldengra-
ben Formation for this assemblage of beds, subdivided
in ‘units’ A, B, C and D. Weidmann (1993) then assigned
the fossil sites to the Staldengraben Formation, more pre-
cisely to the Unit A. e exceptional character of these
beds (i.e. good preservation and abundance of fossils and
organic matter content) seems to be a consequence of the
Toarcian oceanic anoxic event (Mettraux, 1988; Mettraux
& Mosar, 1989). Plancherel etal. (2020) later proposed to
use the term Staldengraben Formation, but to redefine
the Members by adding names of localities where the cor-
responding lithologies are particularly well represented
(proposed type localities). us, the layer, from which
the coelacanth was collected, is assigned to the Soladier
Member of the Staldengraben Formation. e Soladier
Member is composed by sediments predominantly clayey
with alternation of dark schistose marls, altered to brown-
beige, and light marly limestones, mottled, in thin banks
(10–30cm). e coelacanth fossil lies on the cleavage sur-
face of a light-grey to brownish marly limestone that is
then characteristic from the Soladier Member.
e sediments of the Creux de l’Ourssection (Fantasia
etal., 2019) and of Les Pueys were deposited in the west-
ern Tethys in a deep and distal part of the Sub-Brian-
çonnais basin bordered to the south by the Briançonnais
micro-continent and to the north by the Alemanic High
(Fig.1B). Hug (1898) dated the Creux de l’Ours bedsand
of Les Pueys to the early Toarcian based on ammonites.
Weidmann (1993) noted that recent collections of ammo-
nites, carefully collected bed by bed, have revealed the
presence of the Elegantulum and Exaratum Subzones
of the Falcifer Zone (Pugin, 1985) dated of the Toarcian.
During the Toarcian, the marine environment has experi-
enced an important anoxic event known worldwide as the
15 Page 4 of 20
C.Ferrante et al.
‘Toarcian oceanic anoxic event’, which is marked by marine
mass extinctions with a global warming, a perturbation of
the carbon cycle and important depositions of organic-
rich sediments (Fantasia et al., 2018). Sedimentological
analyses in the Creux de l’Ourssection show high kaolinite
contents and detrital proxies suggesting warm and humid
climate during the ‘Toarcian oceanic anoxic event’ (Fanta-
sia etal., 2018). e sediments of the Creux de l’Ours sec-
tion(Fantasia etal., 2018) and of Les Pueys are composed
of grey thin-bedded hemipelagic marls and marly lime-
stones with carbonate concretions formed around accu-
mulations of ostracods and gastropods shells.
Systematic paleontology
Sarcopterygii Romer 1955
Actinistia Cope 1871
Latimerioidei sensu Toriño etal., 2021
Latimeriidae sensu Toriño etal., 2021
Libys Münster 1842
Diagnosis (emended from Forey, 1998)
A genus of medium-sized, relatively deep-bodied coela-
canths. e head is nearly as deep as long. e postpari-
etal shield is much expanded posterolaterally where the
supratemporals are particularly large. e palate is deep
and tapers rapidly anteriorly and the symplectic is also
long, in keeping with the rather deep head. e cheek is
covered with large thin bones which are, however, well
separated from one another. e postorbital and lachry-
mojugal are broad and the preopercle is long and strap-
like. e opercle is substantially deeper than broad. e
subopercle and the preorbital are absent. Sclerotic ossi-
cles are present. In the lower jaw the angular shows a
prominent dorsal expansion and the principal coronoid
Fig. 2 Ammonites and bivalves of Les Pueys compared to ammonites and bivalves of the Creux de l’Ours. Invertebrate fossils A–C preserved on
the reverse side of the two slabs with Libys callolepis sp. nov. (holotype, NMBE 5034072 and 5034073) from Les Pueys and D–F invertebrate fossils
from the Creux de l’Ours. A Dactylioceras? sp. (NMBE 5034073); B Harpoceras renevieri (NMBE 5034072); C Goniomya rhombifera (NMBE 5034072); D
Dactylioceras commune Sowerby 1815 (NMBE 5014840); E Harpoceras renevieri (holotype, NMBE 5014830); F Goniomya rhombifera (NMBE 5021902)
Page 5 of 20 15 The rst Jurassic coelacanth from Switzerland
is developed posterodorsally as a rounded finger-like
process. e most obvious specialization of Libys are the
sensory canals which open to the surface through a large
groove crossed by pillars. Ornamentation is absent from
the skull bones. e shoulder girdle shows a very narrow
cleithrum, clavicle and extracleithrum. e anocleithrum
is simple and sigmoid to blade-like. e pelvic fin is
located well behind the level of the anterior dorsal fin
and is supported by very narrow pelvic bones. e rays of
the anterior dorsal fin and the caudal fin are ornamented
with many prominent denticles. Fin rays of pectoral, ven-
tral, posterior dorsal and anal fins are very closely articu-
lated close to their bases. e supplementary caudal fin is
prominent developing apart from the caudal fin profile.
e lateral line scales carry a large sensory tube which
opens through several secondary tubules. An ossified
lung is present.
L. polypterus Münster 1842
Diagnosis
Libys species with the postparietal shield less than half
the length of the parietonasal shield. e teeth covering
the palate and the lower jaw are very small, and most
are rounded and bear delicate radiating ridges. 70 neu-
ral arches. Fin rays of the posterior dorsal, pectoral,
pelvic and anal fins are expanded. e pectoral fin is
relatively long, reaching back to posterior level of pelvic
fin. e scales are covered with a sparse ornament of
short ridges.
Measurements and meristic
(SL) Standard length 600mm.
d1.f = 10; d2.f = 15–20; pect.f = 16; pelv.f = 19; ana.f = 18–20;
cau.f = 21/19; n.a = 70; h.a = 23.
Holotype
BSM 1870.xrv.502, head only.
Horizon and type locality
Tithonian (Upper Jurassic), Bavaria, Germany.
L. callolepis sp. nov.
Diagnosis
Libys species with the postparietal shield about half the
length of the parietonasal shield (the parietonasal is then
proportionally shorter than in the type species). e teeth
covering the prearticular are very small, and rounded and
smooth. Between 41–47 neural arches. Fin rays are slen-
der than in the type species and then not expanded. e
scales are strongly ornamented with irregularly sized and
elongated round-to-ovoid ridges disposed along a longi-
tudinal axis.
Measurements and meristic
(TB) Total body length 290mm (estimation); (SL) stand-
ard length 255mm.
d1.f = 10;d2.f = 16; pect.f = 18–22;pelv.f ≥ 17;ana.f = 20–23;
cau.f = 15/14–16;n.a = 44–47;h.a = 18–20.
Etymology
From the ancient Greek καλός, kalós, (‘beautiful’,
‘nice’) and λεπίς, lepís, (‘scale’) in reference to the
nicely ornamented scales of the species, which differ-
entiates it from the type species.
Holotype and only known specimen
NMBE 5034072 and 5034073, a sub-complete specimen
preserved in right lateral view as part and counterpart.
Most of the bones, including the scales on the body, are
preserved in anatomical position and only the bones of
the cheek and the jaw are missing. e specimen is kept
in the collections of the Natural History Museum Bern
(Canton of Bern, Switzerland).
Horizon and type locality
Toarcian (Lower Jurassic), Creux de l’Ours section, local-
ity of Les Pueys near the Teysachaux summit (Canton of
Fribourg, Switzerland).
Nomenclatural act
e present work and its nomenclatural act are regis-
tered in ZooBank, the online registration system for
the International Commission on Zoological Nomen-
clature. e Life Science Identifiers for this publication
is “urn:lsid:zoobank.org:pub:D03A2AC9-51F9-45E6-
8CAC-F06A1526AF2C” and can be resolved append-
ing the prefix “http:// zooba nk. org/” in any standard web
browser.
Description
Generalities
Most of the bones of the specimen are preserved on the
part (NMBE 5034073) (Fig.3). From the head, most of the
dermal bones of the skull roof are preserved. e bones of
the cheek and the lower jaw are missing, revealing then the
bones of the palatoquadrate and of the branchial appara-
tus. e axial skeleton, the girdles and the fins are almost
entirely preserved, except the posterior tip of the supple-
mentary caudal fin lobe. e scales are well preserved on
the entire body in natural position, especially on the coun-
terpart (NMBE 5034072) (Fig.4).
15 Page 6 of 20
C.Ferrante et al.
e specimen may represent an adult individual as all the
basal plates are fully ossified, which is a feature observed
in adult coelacanths (e.g., Schultze, 1980; Witzmann etal.,
2010). is view is reinforced because the specimen shows
some well ossified axial mesomere and the scapulocoracoid
(Fig.4A1, A5, A6, B). Although a long supplementary lobe
of the caudal fin is considered as a juvenile character (e.g.,
Forey, 1981; Schultze, 1972), the prominent supplementary
lobe of the caudal fin (Fig.3) observed in the specimen of
Teysachaux represents rather a generic character of the
specimen.
Dermal bones oftheskullroof
e bones of the skull are only preserved on the part
(Figs.3 and 5). e skull roof is divided into a parietona-
sal and postparietal shields, free from one to the other and
separated by the intracranial joint, which appears to be
straight. e postparietal shield appears to be smaller than
the parietonasal shield, this last being about 1.45 longer.
Although slightly lower, this ratio is however close to the
ratios of Jurassic and Cretaceous coelacanths that have typ-
ically a parietonasal shield circa 1.5 to 2 times longer than
the postparietal shield (e.g., Libys polypterus has a ratio of
about 1.7). On the Teysachaux specimen, the dermal bones
of the skull appear to be smooth and unornamented.
Parietonasal shield
e tip of the snout (Fig.3A4, B and Fig.5) is developed
as a heavily ossified hemisphere (Ros.Pmx), which it is not
sutured to the neighboring lateral rostral and rostral ossi-
cles or nasals. It is difficult to discern the premaxilla in this
structure, but the presence of this bone can be deduced
because of a posterior notch that is interpreted here as the
anterior opening for the rostral organ (a.ros). No teeth are
observed, but this could be due to taphonomic processes.
is consolidated snout is reminiscent of the ossified hemi-
spherical snout of Macropoma lewesiensis (Forey, 1998,
figs. 3.19 and 3.20). e difference is that this consoli-
dated snout presents no ornaments, while M. lewesiensis
has a consolidated snout strongly ornamented with small
rounded tubercles (Forey, 1998).
An elongated bone posterior and ventral to the consoli-
dated snout is interpreted as a lateral rostral (L.r) (Figs.3
and 5). is bone has been compressed inside the skull
making the ventral process to point laterally.
Posterior to the consolidated snout lie at least three
nasals (Na) (Figs.3 and 5). e nasal in the middle of the
series is crushed and broken. It is unclear if the sheets of
bone that lie between the anterior nasal and the con-
solidated snout are small rostral ossicles, or a fragmented
nasal. e posterior two nasals are of the same width than
the parietals, and only the anterior most nasal is slightly
narrower. It is difficult to assess the length of the nasals
because of their preservation, but it seems that they are all
of the same length and considerably shorter than the pari-
etals. e lateral margin of the posterior most nasal has
elongated bony pillars that separate large openings of the
supraorbital sensory canal (so.s.c).
e right anterior parietal (Pa) is followed posteriorly by
the right posterior parietal (Figs.3 and 5). e right ante-
rior parietal has its posterior margin crushed. e anterior
portion of the posterior parietal is covered by sediments
and crushed bone, making difficult to assess its length.
However, regarding the length of the anterior parietal, it is
likely that both parietals have the same length. Below both
parietals is a long bone of the left side preserved in inter-
nal view. It is unclear if this bone represents a single bone
(a parietal) with a fracture in its middle or two bones (two
parietals or one parietal plus a nasal). Regarding the present
elements, it appears that both parietals have almost the
same width. Unfortunately, it is unclear if the descending
process of the parietal is present or not. e lateral mar-
gins of the parietals produce thin bony pillars, which are
much more elongated on the anterior parietal (Figs.3A3,
B and 5). ese pillars, when in contact with the neigh-
boring supraorbitals, delimitate very large openings of the
supraorbital sensory canal (so.s.c).
From the lateral series, four posterior supraorbitals (So)
can be clearly observed near the lateral margin of the pos-
terior parietal (Figs.3 and 5). Compared to the length of
the parietals, it is clear that there were more supraorbitals
composing the lateral series. Only one supraorbital lies
directly in contact with the posterior parietal while one or
two posterior most elements are not in direct contact. e
supraorbitals have their mesial margin producing bony bars
or pillars that directly contacts their antimeres on the pari-
etals (Figs.3A3 and B and 5). e openings of the supraor-
bital sensory canal are then proportionally large, almost
forming a groove crossed by pillars. A supraorbital sensory
canal opening through a continuous groove crossed by pil-
lars is reminiscent of Libys polypterus and Megalocoela-
canthus dobiei (Dutel etal., 2012). However, compared to
the two latter taxa, the groove/openings in the specimen of
Teysachaux is a bit less wide. e lateral bony portion of
(See figure on next page.)
Fig. 3 Skeleton of Libys callolepis sp. nov. on the part (holotype, NMBE 5034073). A Photos with osteological details: 1, denticles on the proximal fin
rays of the caudal fin. 2, Postparietal shield with the otic sensory canal opening as a deep groove crossed by pillars (white arrowhead). 3, Posterior
parietal and the supraorbitals with their pillars (white arrowhead). 4, Consolidated snout with the anterior opening for the rostral organ (white
arrowhead). 5, Teeth on the prearticular. B Semi‑interpretative line drawing of the specimen
Page 7 of 20 15 The rst Jurassic coelacanth from Switzerland
Fig. 3 (See legend on previous page.)
15 Page 8 of 20
C.Ferrante et al.
the supraorbitals are narrower compared to the parietals,
but with the pillars the entire bone is almost as large as the
parietals. Regarding the way that the sensory canal opens,
it probably follows a sutural course between the parietals
and the supraorbitals as in most coelacanths.
Apart from these four supraorbitals, two elements rep-
resenting tectals (Te) can be observed in the snout (Figs.3
and 5). ese small bony plates are of the same width than
the supraorbitals. Due to the poor preservation of the lat-
eral series, it is then impossible to determine the exact
number of elements within the series with accuracy.
No preorbital is preserved on the specimen and its pres-
ence/absence remains unknown.
Postparietal shield
e postparietal (Pp) appears to be of the same width than
the posterior parietal and supraorbitals together (Figs.3 and
5). Below the right postparietal extends a large ventral pro-
cess (v.pr.Pp). Unfortunately, the state of preservation makes
impossible to observe if anterior branches of the supratem-
poral commissure, the median branch of the otic canal and
pit lines of the postparietals are marking the bone.
e supratemporal (Stt) is half as long as the postpari-
etal (Figs.3 and 5). e supratemporal bears a large ven-
tral process (v.pr.Stt) stouter than the ventral process of the
postparietal. e otic sensory canal (ot.s.c) separates the
postparietal from the supratemporal as a deep groove but
meet together by thin bony pillars lying within the groove
(Figs.3A2, B and 5).
e supratemporal exceeds the posterior limit of the
postparietal and enclosed a badly crushed bone that
is interpreted here as a lateral extrascapular (Ext.l)
(Figs.3 and 5). Posterior to this bone lies a bone vis-
ible in internal view that could correspond to another
extrascapular. e state of preservation makes difficult
to assess if the extrascapulars are sutured or are free
from the postparietals. e size of the extrascapular
suggests that there were few bones in the extrascapular
series, which is a condition similar to Libys polypterus
and unlike Macropoma (Forey, 1998). e extrascapu-
lar being situated behind the level of the neurocranium
indicates that the braincase has not fused with the
extrascapulars.
All the dermal bones of the skull roof are com-
pletely smooth. e situation is then similar to Libys
polypterus, to the contrary of Macropoma that has the
dermal bones of the skull covered with small round
tubercles (Forey, 1998). e parietals and postparietal
bear no raised areas.
Dermal bones ofthecheek
No cheek bones are preserved. Only a very badly pre-
served opercle (Op) is present (Figs.3–5). Although the
general shape is difficult to assess, the imprint and the
remaining bone show that the opercle is pinched-out
ventrally as in other actinistians as for instance in Libys
polypterus or Macropoma lewesiensis. e preserved por-
tions of the opercle on the part present no ornamenta-
tion and are smooth. On the counterpart, anteriorly and
below the opercle is a poorly preserved imprint of small
scale-like bone with lines of growth, which may possibly
correspond to a bone of the cheek (ch.bo?), such as for
instance the preopercle or subopercle (Fig.4).
No sclerotic ossicles are observed in the specimen of
Teysachaux but the state of preservation precludes to
assess a true absence. In his emended diagnosis of Libys,
Forey (1998) mentioned that sclerotic ossicles are absent
but scored those bones as present. However, in the holo-
type of L. superbus, Reis (1888, pl. 2. fig.2) figured 6 small
rectangular sclerotic ossicles. In the Bamberg Museum
(State of Bavaria, Germany), there is a specimen (NKMB-
P-Watt 08/212) identified as an Undina penicillata that is
in fact a specimen of Libys sp. according to the presence
of a continuous groove crossed by pillars for the supraor-
bital sensory canal and many other anatomical charac-
teristics. is undescribed specimen figured by Mäuser
(2018, fig. 3.2) shows well preserved sclerotic ossicles.
Interestingly, this specimen comes from the Wattendorf
Plattenkalk (State of Bavaria, Germany) that is dated of
the Upper Kimmeridgian (Fürsich etal., 2007) unlike the
holotype of Libys polypterus that is Tithonian in age (e.g.,
Forey, 1998).
Lower jaw
Most of the bones of the lower jaw are not preserved.
Only the left prearticular (Part) is partially visible in
mesial view. Its surface is entirely covered with tiny round
teeth all of the same size arranged in a regular shagreen.
e surface of the teeth appears to be smooth (Fig.3A5).
Fig. 4 Skeleton of Libys callolepis sp. nov. on the counterpart (holotype, NMBE 5034072). A Photos with osteological details: 1, articular head of the
scapulocoracoid. 2, Scales on the flank immediately beneath the first anterior dorsal fin. 3, Scales of the lateral line showing the ornamental pattern
with the larger central tubercles (white arrowheads point, showed only on one scale). 4, Scales on the ventral flank from the pelvic to the anal fin. 5,
Axial mesomere (white arrowhead) surrounded by some fin rays of the anal fin. 6, Axial mesomeres (white arrowhead) partially covered by sediment
in the pelvic fin. B Semi‑interpretative line drawing of the specimen
(See figure on next page.)
Page 9 of 20 15 The rst Jurassic coelacanth from Switzerland
Fig. 4 (See legend on previous page.)
15 Page 10 of 20
C.Ferrante et al.
Fig. 5 Skull of Libys callolepis sp. nov. on the part (holotype, NMBE 5034073). A Photos and B outline of the skull
Page 11 of 20 15 The rst Jurassic coelacanth from Switzerland
Above the prearticular is a fragmented bone that could
possibly belong to the left angular visible in mesial view.
A gular plate (Gu) is visible as a partial and poorly pre-
served imprint.
Neurocranium, palatoquadrate, parasphenoid andgill
arches
Because the right lower jaw is missing, the parasphenoid
and the palatoquadrate can be observed.
e basisphenoid (Bsph) is almost covered by the pala-
toquadrate (Figs.3 and 5). On the visible anterior por-
tion of the basisphenoid, there are two foramens lying
one above the other. e upper foramen, larger than the
lower one, corresponds probably to the opening for the
superficial ophthalmic nerve and the trochlear nerve
(s.oph + IV) while the foramen lying anteroventrally cor-
respond to the foramen for the oculomotor nerve (III).
Just above the metapterygoid, a strong antotic process
(ant.pr) of the basisphenoid is visible.
Posterior to the metapterygoid are preserved fragments
of bone that may correspond to the otic shelf (ot.sh?) of
the prootic, from which can be distinguished a long bony
portion extending posteriorly that could represent the
posterior wing of this bone (Figs.3 and 5). However, this
area is so crushed that it is impossible to discern clearly
the different bones of this region.
Only the anterior half of the parasphenoid (Par) can
be observed, its posterior half being covered by sheet of
indeterminate bone (Figs.3 and 5). Indeed, it seems that
the bone extends posteriorly under the bone interpreted
as the basisphenoid. e main visible anterior portion
bears small tooth. Due to the mode of preservation, it is
hard to assess if the toothed area is restricted to the ante-
rior half or extends more posteriorly. Anteriorly, the par-
asphenoid expands and is upturned dorsally indicating
the presence of a prominent lateral wing (a.w.Par), as for
instance in Macropoma and Latimeria (Forey, 1998). e
contact between the parasphenoid and the basisphenoid,
and then the contact with the processus connectens, can-
not be observed because a sheet of indeterminate bone
lies upon this area. It is probable that the buccohypophy-
sial canal is closed, but this characteristic is difficult to
assert with accuracy.
e right palatoquadrate is well preserved and con-
sists of a pterygoid (Pt) with a quadrate (Q) posteroven-
trally and a metapterygoid (Mpt) dorsally (Figs. 3 and
5). e pterygoid is triangular being as long as it is high
and similar to the palatoquadrate of Macropoma lew-
esiensis (Forey, 1998, Fig.7.2) and Libys polypterus. e
ventral swelling of the pterygoid is well defined as in for
instance Macropoma lewesiensis, Libys polypterus or
Megalocoelacanthus dobiei (Dutel etal., 2012). e exact
shape of the dorsal portion of the metapterygoid is diffi-
cult to clearly assess due to its state of preservation.
e palatoquadrate covers the branchial elements (Cb)
that can nevertheless be partly observed posteriorly
(Figs.3 and 5). e exact number of these elements is dif-
ficult to assess. Some tooth plates (t.p) with tiny granular
teeth accompanied by one or two sharp teeth are pre-
served on some of the branchial elements.
Axial skeleton
e vertebral column is composed of 44 to 47 neural
arches (n.a) and 18 to 20 haemal arches (h.a) (Figs.3 and
4). At least 38 neural spines can be clearly observed from
the tail to the anterior end of the basal plate of the ante-
rior dorsal fin and 3 from this portion to the back of the
skull. In this last portion, there is a pack of superimposed
and badly preserved arches of which the number is hard
to count but should include between 3 to 6 arches. ose
arches lying immediately behind the head (Fig.3) appear
to be broader than the posterior neural arches, being
then expanded according to Forey’s (1998) criteria. ere
are 16 neural arches in the caudal region. Neural and
haemal arches are well spaced and are not abutting. With
a total of 44 to 47 neural arches, the specimen of Tey-
sachaux has less neural arches than in other coelacanths
as for instance Libys polypterus with 70 neural arches,
including 23 haemal arches, and Macropoma lewesiensis
with 60 neural arches, including 22 haemal arches (Forey,
1998).
ere is a maximum of 22 short ossified ribs (ab.rib)
counted on the counterpart (NMBE 5034072), meaning
that they were at least 11 pairs (Figs.3 and 4). Accord-
ing to Lambers (1992) there are 17 pairs of ribs in Libys
polypterus (‘L. superbus’). ose short ossified ribs were
probably confined to the posterior third of the abdomi-
nal area. In many coelacanths, short ribs are confined in
the posterior third of the abdominal area but can be also
present in the anterior half of the abdominal region as in
Holophagus gulo (Forey, 1998, fig.11.8). However, these
short ribs should be distinguished from the long ossified
abdominal ribs observed in the thoracic area of other
actinistians, more generally in mawsoniid coelacanths,
as for instance Diplurus or Chinlea (Cupello etal., 2017;
Forey, 1998).
On the part, in the abdominal region behind the pec-
toral girdle and below the basal plate of the anterior dor-
sal fin, are large fragments of bone that represent a large
ossified lung (oss.lun) (Figs.3 and 4). An ossified lung is
present in other coelacanths as for instance Macropoma,
Libys and Undina, among others (Forey, 1998).
15 Page 12 of 20
C.Ferrante et al.
Paired ns
e basal plates and the girdles are all ossified and are
well preserved. e fin rays of the paired and posterior
unpaired fins are slender than those of the anterior dorsal
fin, being then not expanded unlike in some other coela-
canth as for instance Libys polypterus (Forey, 1998).
Pectoral girdle andns
All bones of the pectoral girdle are present although
badly preserved as imprints and fragmented bones on the
part and counterpart, respectively (Figs.3, 4, 5). All those
bones are smooth and are devoid of ornamentation.
e cleithrum (Cl) is long and narrow throughout its
entire length, and not expanded dorsally. e clavicle
(Cla) is small and tapers anteriorly. e extracleithrum
(Ecl) is a small and elongated ovoid bone. Dorsal to the
cleithrum is a blade-like shaped anocleithrum (Acl).
Forey (1998) mentioned in his emended diagnosis of
Libys polypterus that the anocleithrum is expanded and
scored it as forked. However, Lambers (1992, fig.1 and
pl. 1) described and illustrated a sigmoid shaped ano-
cleithrum on a specimen of Libys polypterus (‘L. super-
bus’). e overall pectoral girdle of the specimen of
Teysachaux does not present any particular anatomi-
cal features, except being particularly narrow as in Libys
polypterus (‘L. superbus’) (Lambers, 1992, fig.1 and pl. 1)
and Undina penicillata (Forey, 1998).
On the counterpart, the left scapulocoracoid (Scc)
is well preserved and extends posteriorly just above
the extracleithrum (Fig.4A1, B). On the part, the right
scapulocoracoid has been shifted from its anatomic
position (Figs.3 and 5). e bone has an articulatory
tip that accommodated the mesomers of the pectoral
fin. It is worth noting that the articulatory tip and the
rest of the bone is entirely ossified, which is remark-
able. Indeed, in Latimeria and probably in most extinct
actinistians, the scapulocoracoid is mostly cartilagi-
nous and only the articulatory tip is ossified (Forey,
1998; Mansuit etal., 2019). e pectoral fin is made of
about 18–22 rays.
Pelvic girdle andn
e pelvic girdle (P.b) and one pelvic fin (pelv.f) are pre-
served on the counterpart (Fig.4). e pelvic girdle lies
in the abdominal area at the same level than the basal
plate of the first dorsal fin. e pelvic fin is then located
well behind the level of anterior dorsal fin as in Libys
(Lambers, 1992, fig.1 and pl. 1; Forey, 1998; Fig.7A).
e pelvic bones are narrow and long with an enlarged
posterior portion similar to that of Libys (Lambers,
1992, fig.1 and pl. 1; Forey, 1998; Fig.7A). As in most
actinistians, both pelvic bones remain separate. It is
difficult to assess clearly the number of fin rays in the
pelvic fin because some of them are preserved one
above the other, and because some rays appear to have
split during the taphonomical process. erefore, as
33 hemi-rays are present, at least 17 fin rays formed in
pelvic fin. In between the fin rays are thin fragments
of bones (Fig.4A6, B) that are interpreted as ossified
axial mesomeres (ax.mes). Although their precise shape
cannot be observed because they are partially covered
by sediments, the outline of the larger one is reminis-
cent of another ossified axial mesomere well preserved
within the anal fin (Fig.4A5, B). It should be noted that
these two pieces of bone could also form a single larger
bone rather than two individual bones.
Unpaired ns
Anterior dorsal n
e basal plate of the anterior dorsal fin (D1.b) is bro-
ken into two parts, the anterior half preserved on the
counterpart and the posterior half on the part (Figs.3
and 4). e basal plate is a rectangular bone strength-
ened in its ventral portion by a ridge running from the
anterior border to the center of the posterior half. e
basal plate lies completely above the level of the neural
spines and has its ventral margin smooth like in most
coelacanths. e anterior dorsal fin (d1.f) is composed
of 10 fin rays (Fig.3). e anterior first ray is compar-
atively shorter than the other rays. e rays are orna-
mented with small and prominent denticles (Fig.3A1).
e number of fin rays is the same than in Libys
polypterus (Forey, 1998). However, the rectangular
shape of the basal plate is different from the triangular
basal plate of Libys polypterus (‘Libys superbus’) (Forey,
1998; Lambers, 1992, fig.1 and pl. 1) and Macropoma
lewesiensis (Forey, 1998 fig.11.11).
Posterior dorsal n
On the part, the basal plate (D2.b) and rays of the pos-
terior dorsal fin (d2.f) are preserved while there are only
some fin rays on the counterpart (Figs.3 and 4). About
16 fin rays are counted on the part and the counterpart.
e rays are devoid of denticles. e basal plate of the
posterior dorsal fin is a bone formed by an ovoid plate
extending forward by two thin rods. e two rods of the
bifurcated anterior portion are of the same length. e
main plate bears on its anterodorsal corner a small bony
process developing anteriorly, which is present in Latim-
eria and Laugia (Millot & Anthony, 1958, pl. LIXa; Forey,
1998, figs. 8.3a and 8.3c) and potentially in Piveteauia
(Clément, 1999).
Page 13 of 20 15 The rst Jurassic coelacanth from Switzerland
Anal n
e anal basal plate (A.b) is preserved in two halves, one
on the part and the other on the counterpart (Figs.3 and
4). e rays of the anal fin (ana.f) are only preserved on
the counterpart and comprise about 20 to 23 rays. e
anal fin is positioned as a mirror image of the second
dorsal fin. e posterior portion of the bone is rectangu-
lar and spatula-shaped widening slightly posteriorly. e
anterior part is bifurcated in two long processes symmet-
rically positioned. ese processes are shorter than the
two processes of the basal plate of the posterior dorsal
fin. It is interesting to note that a small axial mesomere
(ax.mes) supporting the fin rays can be observed on the
counterpart (Fig.4A5). Axial mesomeres are rarely pre-
served because the endoskeleton is usually cartilaginous.
e Early Triassic Laugia is one of the rare cases where
those bones are ossified (Forey, 1998).
Caudal n andsupplementary lobe
e caudal fin (cau.f) is preserved on the part and pre-
sent the diphycercal pattern with a lower and upper lobes
separated by a supplementary caudal lobe typical of coe-
lacanths (Fig.3). e caudal fin contains 16 and about
14 radials (Ra) in the upper and lower lobe, respectively.
ese bones have proximal and distal expanded extremi-
ties. In the upper and probably also in the lower lobe,
the first anterior radial doesn’t bear any fin ray. ere
are 15 and 14 to 16 fin rays in the upper and lower lobes,
respectively. According to the definition of Forey (1998),
the tail is symmetric. In the upper and the lower lobe,
respectively, the sixth and fourth anterior most rays bear
sharp and prominent denticles while posteriorly the rays
are devoid of any denticles (Fig.3A1).
e supplementary caudal lobe (sup.cau.f.l) is present
but the tip is unfortunately missing (Fig. 3). Although
incomplete, the supplementary caudal lobe is compara-
tively long and prominent appearing to develop apart
from the caudal fin profile, which is reminiscent of Libys
polypterus (‘L. superbus’) (Lambers, 1992, fig. 1 and pl.
1B-C) and Undina penicillata (Forey, 1998; Fig.7). is
feature is considered in the specimen of Teysachaux, as
well as in the genus Libys in general, as a generic charac-
ter rather than the indication of a juvenile stage found in
some other examples of coelacanths (e.g., Schultze, 1972).
Indeed, in the examined specimens of Libys with the sup-
plementary caudal lobe kept in the BSM, the basal plate
is always fully ossified indicating an adult state. ere-
fore, these coelacanths have a supplementary caudal lobe
developing well apart from the caudal fin unlike other
coelacanths that have a supplementary lobe enclosed in
the caudal fin such as for instance Macropomoides (Forey,
1998) or Foreyia (Cavin etal., 2017, fig.1).
Scales
e scales are well preserved in natural position and
exhibit their ornamental pattern, which shows varia-
tions according to their position on the body (Fig.4A2).
e exact shape of the entire scales cannot be assessed
and only the exposed ornamented part, which is triangu-
lar, can be observed. On the flank immediately beneath
the first anterior dorsal fin, the scales are strongly orna-
mented with irregularly sized and elongated round-to-
ovoid ridges disposed along a longitudinal axis (Fig.4A2).
Each scale of the lateral line bears three round and
pointed tubercles in its middle exposed area that are sur-
rounded by short to elongated ovoid ridges (Fig. 4A3).
e pores of the lateral line are very difficult to observe
but on some very rare scales, multiple small pores sur-
rounding the central tubercles can be recognized. On the
ventral flank, from the pelvic to the anal fins, the scales
are ornamented with delicate elongated ridges (Fig.4A4).
No ventral keel scales are present. e patterns of the
scales and their distribution on the body of the specimen
of Teysachaux are reminiscent of the scales of Macro-
poma lewesiensis as figured by Woodward (1909, pl. 38.
fig.4a–c) and Forey (1998, fig.11.12a–b). ey are dif-
ferent from Libys polypterus (Fig.6B), Undina penicillata
(Fig.6C) and Trachymetopon (‘Macropoma’) substrialo-
tum (Fig.6D), in which the scales are ornamented with
dense to sparse ornament of thinner tubercles.
Discussion
Identication andcomparison oftheTeysachaux specimen
withother Mesozoic coelacanths
When the specimen was integrated in 1873–1874 within
the collection of the Museum of Bern, Fischer-Ooster
gave it the name of ‘Macropoma heeri’. However, Fis-
cher-Ooster never properly published this species with a
description and illustration, probably because he passed
away in 1875. Since its discovery, the specimen was only
cited under the name ‘Macropoma’, first by Hug (1898)
in his description of the fauna of Les Pueys and, then by
Marchant and Pichon (2013), who illustrated it by a pho-
tograph in the guide ‘Jurassique Suisse’. Consequently,
because the specific name has not been published with
an adequate scientific description, and according to the
International Code of Zoological Nomenclature rules,
the species name ‘heeri’ is considered as nomen nudum.
In 1873–1874, only few genera of coelacanths were
known, i.e. Macropoma, Undina, Coelacanthus, Libys,
Rhabdoderma, Heptanema, Coccoderma, Holophagus
and Graphiurichthys. us, it is probable that Fischer-
Ooster gave the generic name Macropoma because
some details, especially the scales, are reminiscent
of Macropoma. e main question is whether the
15 Page 14 of 20
C.Ferrante et al.
coelacanth of Teysachaux belongs to a currently known
genus and species or represents a new taxon.
e specimen of Teysachaux presents the following
combination of characters. One of its most remark-
able morphological features is its sensory canal opening
though a large groove crossed by pillars. is peculiar
characteristic is currently known in Libys (Lambers,
1992, fig.7 pl. 1B-C; Fig.7A) from the Upper Jurassic
of Germany (Forey, 1998) and in Megalocoelacanthus
from the Upper Cretaceous of the United States (Dutel
etal., 2012, figs.2, 3, 5 and 7). e specimen of Tey-
sachaux is then different from Macropoma because
in all species of this genus the sensory canals open
through many small pores (Forey, 1998).
e consolidated snout of the specimen of Teysachaux
is a particular morphological feature found in few coela-
canth taxa as Laugia groenlandica (Stensiö, 1932, pl. 5.
figs.4–5), Macropoma lewesiensis (Forey, 1998, figs.3.19a
and 3.20), Swenzia latimerae (Clément, 2005, fig.4) and
Megalocoelacanthus dobiei (Dutel et al., 2012, fig. 4).
Compared with these taxa, the consolidated snout of the
specimen of Teysachaux (Figs.3A4 and 5) is rather simi-
lar to that of M. lewesiensis having an anterior opening of
the rostral organ occurring as a notch between the ossi-
fied snout and the rostral ossicles (Forey, 1998, figs.3.19a
and 3.20a). However, conversely to M. lewesiensis the
snout of the specimen of Teysachaux is not ornamented.
Unfortunately, it is not possible to make a reasonable
Fig. 6 Comparison of scales between Libys callolepis sp. nov. and some other Latimeriidae. A Libys callolepis sp. nov. (holotype, NMBE 5034072).
Scales of the flank below the neural arches between the basal plates of the anterior and posterior dorsal fin. B Libys polypterus (‘L. superbus’) (BSM AS
I 801a). Scales of the flank located between the neural arches and the basal plate of the second dorsal fin. C Undina penicillata (BSM 1870 XIV 22).
Scales of the middle flank located on the neural arches below the basal plate of the second dorsal fin. D Trachymetopon (‘Macropoma’) substriolatum
(holotype, SMC J27415) from the Kimmeridgian of Cottenham, Cambridgeshire. Scales of the middle flank. It is worth noting that this species was
first attributed by Huxley (1866) to the genus Macropoma and latter to Coccoderma by Reis (1888). Forey (1998) kept this species within Coccoderma,
but remarked that it shows affinities with Holophagus. Recently, Cavin et al. (2021a) attributed this specimen to the genus of Trachymetopon
regarding, among other characteristics, the coarse ornamentation on the dermal bones
Page 15 of 20 15 The rst Jurassic coelacanth from Switzerland
comparison with the snout of L. polypterus because
this portion is still imperfectly known and remain to be
described on well preserved specimens.
Regarding the meristic features, the specimen of Tey-
sachaux is more closely related to Libys polypterus than
to Macropoma lewesiensis, Undina penicillata or Holo-
phagus gulo (Table1). e specimen of Teysachaux and
L. polypterus have both 10 rays in the anterior dorsal fin,
while H. gulo has 10–11 rays and M. lewesiensis and U.
penicillata have both less rays with 7 and 8 rays, respec-
tively (Forey, 1998). Although sharing similarities with L.
polypterus, the specimen of Teysachaux has 44 to 47 neu-
ral arches in the vertebral column, which is less than in
M. lewesiensis, H. gulo, L. polypterus and U. penicillata
Fig. 7 Photographs of two latimeriid coelacanths from the Upper Jurassic of Germany showing the body outline and caudal fin with
a prominent supplementary caudal lobe. A Specimen of Libys sp. (no collection number) exposed in the permanent exhibition of the
Bürgermeister‑Müller‑Museum, Solnhofen, Germany. It is worth noting that this specimen is labeled as Holophagus, but the morphological
characteristics (e.g., a sensory canal opening through a large groove crossed by pillars) and the meristic clearly indicate that this specimen belongs
to the genus Libys. B Undina penicillata (BSM 1870 XIV 22)
15 Page 16 of 20
C.Ferrante et al.
that have 60, 65, 70 and 70–72 neural arches, respec-
tively (Forey, 1998). Forey (1998) remarked that there is
considerable variation among coelacanths regarding the
number and the relative spacing of the neural arches.
According to Forey (1998), plesiomorphic coelacanths
have fewer neural arches than derived taxa. While not-
ing that there are some exceptions, Forey (1998) gave the
example of Rhabdoderma with 45 neural arches as repre-
senting a plesiomorphic state and Latimeria with about
110 neural arches. erefore, compared to L. polypterus,
M. lewesiensis, H. gulo and U. penicillata, the specimen
of Teysachaux represent a more plesiomorphic state.
e prominent supplementary caudal lobe of the
specimen of Teysachaux is clearly reminiscent of the
prominent supplementary caudal lobe of the genus Libys
(Lambers, 1992, fig.1, pls 1A-B and 2A; Fig.7A). e sup-
plementary caudal lobe of Undina penicillata (Fig.7B) is
also long (Forey, 1998) but much less prominent than in
the specimen of Teysachaux and L. polypterus. Unfortu-
nately, the supplementary caudal lobe of Macropoma is
unknown (Forey, 1998).
e ornamentation of the scales of the specimen of
Teysachaux is reminiscent of the ornamentation of the
scales of Macropoma lewesiensis (Forey, 1998, fig.11.12
a-b) being then different from Libys polypterus (Fig.6B).
Furthermore, the variation of the type of ornamenta-
tion according to the position on the body is also simi-
lar in the specimen of Teysachaux (Fig.4A3-4A4) and in
Macropoma lewesiensis (Forey, 1998, figs.11.12 a-b).
It is also important to make some comparative remarks
with Undina (?) barroviensis, which is a poorly defined
species of uncertain affinities described first by Wood-
ward (1890) from the Lower Jurassic of England. Accord-
ing toForey (1998) Undina (?) barroviensis has relatively
large supraorbital sensory pores. Forey (1998) used the
term "very large pores" to describe the openings of the
sensory canal of Libys polypterus. It therefore remains
to demonstrate that this character described by Forey
(1998) is the same as in L. polypterus. Among other
known characters, Forey (1998) also noticed that in
Undina (?) barroviensis each scale bears few stout tuber-
cles. e current state of knowledge concerning Undina
(?) barroviensis is insufficient to make a more relevant
comparison with the specimen of Teysachaux. However,
Undina (?) barroviensis and the specimen of Teysachaux
clearly belongs to different species because the number
of neural arches in the first species is significantly larger
than in the second one, i.e. more than 50 based on a
comparison with Holophagus gulo made by Forey (1998).
According to the previous remarks, the specimen
of Teysachaux shares many characters with the genus
Libys, i.e. (1) head nearly as deep as long; (2) openings
of the sensory canals through large grooves crossed by
pillars; (3) dermal bones of the skull smooth and unor-
namented; (4) deep palatoquadrate with a pronounced
ventral swelling; (5) very narrow cleithrum, clavicle and
extracleithrum; (6) pelvic fins located well behind the
level of the anterior dorsal fin and supported by narrow
pelvic bones; (7) same number of rays in the anterior
and posterior dorsal fins and in the anal fin; (8) promi-
nent supplementary caudal lobe; (9) rays of the anterior
dorsal fin and the caudal fin ornamented with many
prominent denticles and; (10) presence of an ossified
lung.
Nevertheless, the specimen of Teysachaux presents
some different morphological characters from Libys
polypterus, such as: (1) a postparietal shield about half the
length of the parietonasal shield (the parietonasal shield
is then proportionally shorter than in L. polypterus);
(2) the teeth covering the prearticular are very small,
rounded and smooth; (3) less neural and haemal arches;
(4) all fin rays are slender and then not expanded and;
(5) scales strongly ornamented with irregularly sized and
elongated round-to-ovoid ridges disposed along a longi-
tudinal axis.
erefore, the specimen of Teysachaux does not belong
to the genus Macropoma as suggested by von Fischer-
Ooster. e specimen of Teysachaux is reminiscent of
Libys polypterus Münster 1842 but based on morpho-
logical differences noted above, it should be regarded as
a new species of Libys. erefore, we placed the specimen
described here in a new species of the genus Libys and
erect the new species L. callolepis sp. nov.
Table 1 Meristic of some Latimeriidae compared to Libys callolepis sp. nov. Dataset from Forey (1998)
Taxon/No. of elements d1.f d2.f pect.f pelv.f ana.f cau.f (up.\lo.lobe) n.a
Latimeria chalumnae 8 29–31 30–32 33 29–32 22–25/21–22 93–94
Macropoma lewesiensis 7 17 16 18 16 18–20/15–18 60
Undina penicillata 8 15–17 19 21 14 18/16 70–72
Holophagus gulo 10–11 20 23 18 17 18/17 65
Libys polypterus 10 15–20 16 19 18–20 21/19 70
Libys calloelpis sp. nov. 10 16 18–22 ≥ 17 20–23 15/14–16 44–47
Page 17 of 20 15 The rst Jurassic coelacanth from Switzerland
Implication fortheevolutionary history ofcoelacanths
In the Tithonian (Upper Jurassic) of the Solnhofen region
(Bavaria, Germany), two species of Libys have originally
been recognized: L. polypterus Münster, 1842, and L. super-
bus Zittel, 1887. However, Forey (1998) and subsequent
authors considered L. superbus to be the junior synonym
of L. polypterus. e presence of a new species of Libys in
the Toarcian of Teysachaux (Canton of Fribourg, Switzer-
land) extends the stratigraphic range of this genus by about
34 million years. is time interval may seem long, but it
is comparable to the estimated time interval for the extant
Latimeria (Inoue etal., 2005; Sudarto etal., 2010) and the
stratigraphic range is still longer for 12 extinct genera of
coelacanths (Cavin etal., 2021b). In the most recent phy-
logenies of coelacanths (e.g., Toriño etal., 2021), Libys is
resolved as the sister genus of the Upper Cretaceous Mega-
locoelacanthus, both branching in the family Latimeriidae.
Although the stratigraphic range of the genus Libys has
expanded considerably, its geographical distribution has
not considerably increased. However, the Middle Jurassic
fossil record of coelacanths, separating the Lower Juras-
sic Swiss occurrence of L. callolepis sp. nov. from the Late
Jurassic German occurrence of L. polypterus, is particularly
poor with only one other genus recorded.
e taxic diversity of the coelacanths estimated on
the basis of the fossil record has always been low since
the Lower Devonian. e diversity shows some vagar-
ies over time with periods of proportionally high and
low diversity (Fig. 8a). However, when calculated per
million years rather than raw diversity, the peak and the
falls are smoothed (Fig.8b). In this pattern, coelacanths
experienced a very high peak in taxa diversity after the
Permo-Triassic mass extinction in the Lower Triassic
and to a lesser degree in the Middle Triassic. From the
beginning of the Upper Triassic, the taxic diversity of the
group shows a significant steady decrease until the Mid-
dle Jurassic where it reaches its minimum with a single
known species. e cause of the decline of coelacanths
may be related to competition with neopterygians and
chondrichthyans due to their slow metabolism. Indeed,
compared to other bony fish groups, Neopterygians
show a continuous increase in diversity from the Middle-
Upper Triassic (Romano etal., 2016).
Fig. 8 Taxic diversity of coelacanths through time. A Number of observed coelacanth genera by Epoch and B Ratio of coelacanth genera by Epoch,
calculated as the number of genera by Epoch per million of years. Dataset after Cavin et al. (2021a), with the addition of the occurrence of Libys in
the Lower Jurassic
15 Page 18 of 20
C.Ferrante et al.
Alongside a peak of diversity of coelacanths (e.g., Cavin
et al., 2013), the Early and Middle Triassic also wit-
nessed a high morphological disparity, exemplified by the
strange Foreyia from the Prosanto Formation (Canton of
Graubünden) (Cavin etal., 2017), and by another mor-
phologically aberrant taxon from the Besano Formation
(Canton of Ticino), under study (Ferrante et al., 2017;
Ferrante etal., workin progress). e appearance of Libys
callolepis sp. nov. from the Lower Jurassic announces the
beginning of a new stage in the evolution of the coela-
canths, with the appearance of taxa with morphological
Bauplan which does not derogate from the general coe-
lacanth Bauplan (except in disparity of body size (Cavin
etal., 2021a)) and with long stratigraphic ranges and/or
ghost ranges (Toriño etal., 2021).
Conclusion
Libys callolepis sp. nov. represents the only Swiss occur-
rence of a coelacanth, with the exception of those from
the rich Middle Triassic sites of the Prosanto Formation
and of Monte San Giorgio. Its presence in the Lower
Jurassic beds extends the stratigraphic range of the genus
Libys by about 34 million years, but without increas-
ing considerably its geographic distribution. e genus
Libys belongs to the modern family Latimeriidae and the
appearance of L. callolepis sp. nov. heralds a long period,
up to the present day, of coelacanth genera with very long
stratigraphic range and reduced morphological disparity,
which have earned them the nickname of ‘living fossils’.
Abbreviations
NMBE: Naturhistorisches Museum Bern, Bern, Switzerland; BSM: Bayerische
Staatssammlung für Paläontologie und Geologie, Munich, Germany; SMC:
Sedgwick Museum, Cambridge University, England; A.b: Basal plate of the
anal fin; a.ros: Anterior opening for the rostral organ; a.w.Par: Ascending
wing of parasphenoid; ab.rib: Abdominal rib; ax.mes: Axial mesomere; Acl:
Anocleithrum; ana.f: Anal fin; ant.pr: Antotic process; Bsph: Basisphenoid; cau.f:
Caudal fin; Cb: Ceratobranchial; ch.bo: Cheek bone; Cl: Cleithrum; Cla: Clavicle;
D1.b: Basal plate of the anterior dorsal fin; D2.b: Basal plate of the posterior
dorsal fin; d1.f: Anterior dorsal fin; d2.f: Posterior dorsal fin; Ecl: Extracleithrum;
Ext.l: Lateral extrascapular; Gu: Gular plate; h.a: Haemal arches; L.r: Lateral
rostral; Mpt: Metapterygoid; n.a: Neural arches; Na: Nasal; Op: Opercle; oss.
lun: Ossified lung; Ot.sh: Otic shelf; ot.s.c: Otic sensory canal; Pa: Parietal; Par:
Parasphenoid; Part: Prearticular; P.b: Pelvic bone; pect.f: Pectoral fin; pelv.f:
Pelvic fin; Pp: Postparietal; Pt: Pterygoid; Q: Quadrate; Ra: Radial; Ros.Pmx:
Rostropremaxilla; s.oph + IV: Opening for the superficial ophthalmic nerve and
the trochlear nerve; Scc: Scapulocoracoid; So: Supraorbital; so.s.c: Supraorbital
sensory canal; Stt: Supratemporal; sup.cau.f.l: Supplementary caudal fin lobe;
t.p: Tooth plate; Te: Tectal; v.pr.Pp: Ventral descending process of the postpari‑
etal; v.pr.stt: Ventral descending process of supratemporal; III: Foramen for the
oculomotor nerve.
Acknowledgements
We would like to thank Bernhard Hostettler for the ’re‑discovery’ and prepara‑
tion of this coelacanth in the collection of the Natural History Museum Bern.
We are grateful to Adriana López‑Arbarello and Baran Karapunar (Bayerische
Staatssammlung für Paläontologie und Geologie, Munich) for the access
to the collections and for determining the bivalve, respectively. We thank
Rossana Martini for supervising the PhD of CF. We thank both anonymous
reviewers for their constructive comments and the editor Daniel Marty.
Author contributions
CF designed the study, wrote the description of the fossil material, photo‑
graphed and prepared the figures. UM‑G clarified the lithostratigraphy and
biostratigraphy of the site and the studied rock slab. LC supervised the work
(which is a part of the PhD) of CF. All the authors contributed to and approved
the drafting of the last version of the text. All authors read and approved the
final manuscript.
Funding
This paper is a contribution to the project ‘Evolutionary pace in the coelacanth
clade: New evidence from the Triassic of Switzerland’ supported by the Swiss
National Science Foundation (200021‑172700).
Availability of data and materials
The holotype (NMBE 5034072 and 5034073) is kept in the collections of the
Natural History Museum Bern (Canton of Bern, Switzerland). The dataset used
to perform the taxic diversity of coelacanths through time can be found as a
supplementary information from the paper of “Cavin, L., Piuz, A., Ferrante, C., &
Guinot, G. (2021a). Giant Mesozoic coelacanths (Osteichthyes, Actinistia) reveal
high body size disparity decoupled from taxic diversity. Scientific reports,
11(1), 1–13. https:// doi. org/ 10. 1038/ s41598‑ 021‑ 90962‑5”.
Declarations
Competing interests
The authors declare that they have no competing interests.
Author details
1 Department of Earth Sciences, University of Geneva, Rue des Maraîchers 13,
1205 Geneva, Switzerland. 2 Natural History Museum Bern, Bernastrasse 15,
3005 Bern, Switzerland. 3 Department of Geology and Palaeontology, Natural
History Museum of Geneva, CP 6434, 1211 Geneva 8, Geneva, Switzerland.
Received: 11 May 2022 Accepted: 18 July 2022
References
Arratia, G., & Schultze, H. P. (2015). A new fossil actinistian from the early
Jurassic of Chile and its bearing on the phylogeny of Actinistia. Journal of
Vertebrate Paleontology. https:// doi. org/ 10. 1080/ 02724 634. 2015. 983524
Cavin, L., Furrer, H., & Obrist, C. (2013). New coelacanth material from the Mid‑
dle Triassic of eastern Switzerland, and comments on the taxic diversity of
actinistans. Swiss Journal of Geosciences, 106(2), 161–177. https:// doi. org/
10. 1007/ s00015‑ 013‑ 0143‑7
Cavin, L., Mennecart, B., Obrist, C., Costeur, L., & Furrer, H. (2017). Hetero‑
chronic evolution explains novel body shape in a Triassic coelacanth
from Switzerland. Scientific Reports, 7(1), 1–7. https:// doi. org/ 10. 1038/
s41598‑ 017‑ 13796‑0
Cavin, L., Piuz, A., Ferrante, C., & Guinot, G. (2021a). Giant Mesozoic coelacanths
(Osteichthyes, Actinistia) reveal high body size disparity decoupled from
taxic diversity. Scientific Reports, 11(1), 1–13. https:// doi. org/ 10. 1038/
s41598‑ 021‑ 90962‑5
Cavin, L., Toriño, P., Van Vranken, N., Carter, B., Polcyn, M. J., & Winkler, D. (2021b).
The first late cretaceous mawsoniid coelacanth (Sarcopterygii: Actinistia)
from North America: evidence of a lineage of extinct ‘living fossils.’ PLoS
ONE. https:// doi. org/ 10. 1371/ journ al. pone. 02592 92
Clément, G. (1999). The actinistian (Sarcopterygii) Piveteauia madagascariensis
Lehman from the lower Triassic of northwestern Madagascar: A rede‑
scription on the basis of new material. Journal of Vertebrate Paleontology,
19(2), 234–242. https:// doi. org/ 10. 1080/ 02724 634. 1999. 10011 137
Clément, G. (2005). A new coelacanth (Actinistia, Sarcopterygii) from the
Jurassic of France, and the question of the closest relative fossil to
Latimeria. Journal of Vertebrate Paleontology, 25(3), 481–491. https:// doi.
org/ 10. 1671/ 0272‑ 4634(2005) 025[0481: ANCASF] 2.0. CO;2
Page 19 of 20 15 The rst Jurassic coelacanth from Switzerland
Cupello, C., Meunier, F. J., Herbin, M., Janvier, P., Clément, G., & Brito, P. M.
(2017). The homology and function of the lung plates in extant and
fossil coelacanths. Scientific Reports, 7(1), 1–8. https:// doi. org/ 10. 1038/
s41598‑ 017‑ 09327‑6
Dutel, H., Herbin, M., & Clément, G. (2015). First occurrence of a mawsoniid
coelacanth in the Early Jurassic of Europe. Journal of Vertebrate Paleon-
tology. https:// doi. org/ 10. 1080/ 02724 634. 2014. 929581
Dutel, H., Maisey, J. G., Schwimmer, D. R., Janvier, P., Herbin, M., & Clément,
G. (2012). The giant Cretaceous Coelacanth (Actinistia, Sarcopterygii)
Megalocoelacanthus dobiei Schwimmer, Stewart and Williams, 1994,
and its bearing on Latimerioidei interrelationships. PLoS ONE. https://
doi. org/ 10. 1371/ journ al. pone. 00499 11
Fantasia, A., Föllmi, K. B., Adatte, T., Spangenberg, J. E., & Montero‑Serrano,
J.‑C. (2018). The early Toarcian oceanic anoxic event: Paleoenvironmen‑
tal and paleoclimatic change across the Alpine Tethys (Switzerland).
Global and Planetary Change, 162, 53–68. https:// doi. org/ 10. 1016/j.
glopl acha. 2018. 01. 008
Fantasia, A., Föllmi, K. B., Adatte, T., Spangenberg, J. E., & Mattioli, E. (2019).
Expression of the Toarcian oceanic anoxic event: New insights from a
Swiss transect. Sedimentology, 66(1), 262–284. https:// doi. org/ 10. 1111/
sed. 12527
Ferrante, C., Martini, R., Furrer, H., & Cavin, L. (2017). Coelacanths from the
Middle Triassic of Switzerland and the pace of actinistian evolution.
Research Knowledge, 3(2), 59–62. https:// doi. org/ 10. 14456/ randk. 2017.
28
Fischer‑Ooster, C., & Ooster, W. A. (1870). von Über Ichtyosaurus tenuirostris
(Conybeare) aus den Liasschichten am westlichen Fusses des Moleson
in den Freiburger‑Alpen. Protozoe Helvetica, 2, 73–84.
Forey, P. L. (1981). The coelacanth Rhabdoderma in the Carboniferous of the
British Isles. Palaeontology London, 24, 203–229.
Forey, P. L. (1998). History of the coelacanth fishes. London: Chapman and
Hall.
Furrer, H. (1960). Der Ichthyosaurus vom Teysachaux. Mitt Natf Ges Bern, N F,
18, 75–82.
Fürsich, F. T., Mäuser, M., Schneider, S., & Werner, W. (2007). The Wattendorf
Plattenkalk (Upper Kimmeridgian) – a new conservation lagerstätte
from the northern Franconian Alb, Southern Germany. Neues Jahrbuch
Für Geologie Und Paläontologie, Abhandlungen, 245, 45–58.
Hug, O. (1898). Beiträge zur Kenntnis der Lias ‑ und Dogger ‑ Ammoniten
aus der Zone der Freiburger Alpen, I. Die Oberlias‑Ammoniten‑Fauna
von Les Pueys und Teysachaux am Moléson. Abhandlungen der Sch-
weizerischen Paläontologischen Gesellschaft, 25, 1–28.
Huxley, T. H. (1866). Illustrations of the structure of the crossopterygian
ganoids. Figures and descriptions illustrative of British organic remains.
Memoirs of the Geological Survey of the United Kingdom, London, Dec, 12,
1–44.
Inoue, J. G., Miya, M., Venkatesh, B., & Nishida, M. (2005). The mitochondrial
genome of Indonesian coelacanth Latimeria menadoensis (Sarcop‑
terygii: Coelacanthiformes) and divergence time estimation between
the two coelacanths. Gene, 349, 227–235. https:// doi. org/ 10. 1016/j.
gene. 2005. 01. 008
Jain, S. L. (1974). Indocoelacanthus robustus n. gen., n. sp. (Coelacanthidae,
Lower Jurassic), the first fossil coelacanth from India. Journal of Paleon-
tology, 48, 49–62.
Lambers, P. (1992). On the Ichthyofauna of the Solnhofen Lithographic
Limestone (Upper Jurassic, Germany). PhD Thesis. pp. 336.
Mansuit, R., Clément, G., Herrel, A., Dutel, H., Tafforeau, P., Santin, M. D., &
Herbin, M. (2019). Development and growth of the pectoral girdle and
fin skeleton in the extant coelacanth Latimeria chalumnae. Journal of
Anatomy, 236(3), 493–509. https:// doi. org/ 10. 1111/ joa. 13115
Marchant, R., & Pichon, B. (2013). Jurassic Suisse. Ed. Favre SA, Lausanne. pp.
231.
Mäuser, M. (2018). BAMBERG: The paleontological collection at the museum
of natural history in Bamberg (NKMB). Paleontological collections of Ger-
many, Austria and Switzerland (pp. 23–26). Cham: Springer.
Menkveld‑Gfeller, U. (1998). Der Fischsaurier vom Teysachaux: Ein echter
Freiburger? Schweizer Strahler, 11(8), 345–347.
Mennecart, B., & Havran, M. (2013). Nouvelles données sur les Ichtyosaures
du Canton de Fribourg. Bulletin de la Société Fribourgeoise des Sciences
Naturelles, 102, 77–84.
Mettraux, M. (1988). Sédimentologie, paléotectonique et paléo‑océanogra‑
phie des Préalpes médianes (Suisse romande) du Rhétien au Toarcien.
Thèse Université de Fribourg, 947.
Mettraux, M., & Mosar, J. (1989). Tectonique alpine et paléotectonique
liasique dans les Préalpes Médianes en rive droite du Rhône. Eclogae
Geologicae Helvetiae, 82(2), 517–540.
Millot, J., & Anthony, J. (1958). Anatomie de Latimeria chalumnae – Tome I:
squelette, muscles et formations de soutien. Vol. 1. Paris: C.N.R.S. pp.
122.
Plancherel, R., Braillard, L., & Dall’Agnolo, S. (2020). Feuille 1245 Château‑
d’Oex. Atlas géol. Suisse 1:25 000. Notice Explicative, 144, 1–123.
Pugin, L. (1985). Le Toarcien inférieur du Creux de l’Ours (Rapport inedit du
30.05.1985). Institut de Géologie de l’Université de Fribourg.
Reis, O. M. (1888). Die Coelacanthinen mit besonderer Berücksichtigung der
im Weissen Jura Bayerns vorkommenden Arten. Palaeontographica,
35(1), 1–96.
Renesto, S., Magnani, F., & Stockar, R. (2021). A new coelacanth specimen
with elongate ribs from the Middle Triassic (Ladinian) Kalkschieferzone
of Monte San Giorgio (Canton Ticino, Switzerland). Rivista Italiana di
Paleontologia e Stratigrafia, 127(3), 689–700. https:// doi. org/ 10. 13130/
2039‑ 4942/ 16731
Renesto, S., & Stockar, R. (2018). First record of a coelacanth fish from the
Middle Triassic Meride Limestone of Monte San Giorgio (Canton Ticino,
Switzerland). Rivista Italiana di Paleontologia e Stratigrafia, 124(3),
639–653. https:// doi. org/ 10. 13130/ 2039‑ 4942/ 10771
Rieppel, O. (1980). A new coelacanth from the Middle Triassic of Monte San
Giorgio, Switzerland. Eclogae Geologicae Helvetiae, 73, 921–939.
Rieppel, O. (1985). A second actinistian from the Middle Triassic of Monte
San Giorgio, Kt. Tessin. Switzerland. Eclogae Geologicae Helvetiae, 78,
707–713.
Romano, C., Koot, M. B., Kogan, I., Brayard, A., Minikh, A. V., Brinkmann, W.,
Bucher, H., & Kriwet, J. (2016). Permian‑Triassic Osteichthyes (bony
fishes): Diversity dynamics and body size evolution. Biological Reviews,
91(1), 106–147. https:// doi. org/ 10. 1111/ brv. 12161
Schaeffer, B. (1948). A study of Diplurus longicaudatus with notes on the
body form and locomotion of the Coelacanthini. American Museum
Novitates, 1378, 1–32.
Schultze, H.‑P. (1972). Early growth stages in coelacanth fishes. Nature New
Biology, 236, 90–91. https:// doi. org/ 10. 1038/ newbi o2360 90a0
Schultze, H.‑P. (1980). Eier legende und lebendgebärende Quastenflosser.
Natur und Museum, 110, 101–108.
Scotese, C.R., (2014). Atlas of Jurassic Paleogeographic Maps, PALEOMAP
Atlas for ArcGIS, vol. 3, The Jurassic and Triassic, Maps 32–42, Mollweide
Projection, PALEOMAP Project, Evanston, IL.
Septfontaine, M. (1983). Le Dogger des Préalpes médianes suisses et
françaises : stratigraphie, évolution paléogéographique et paléotec‑
tonique. Mémoires de la Société helvétique des sciences naturelles, 97,
1–121. https:// archi ve‑ ouver te. unige. ch/ unige: 149604
Stensiö, E. A. (1932). Triassic fishes from East Greenland. Meddelelser Om
Grønland, Copenhagen, 83, 1–305.
Sudarto, Lalu, X. C., Kosen, J. D., et al. (2010). Mitochondrial genomic
divergence in coelacanths (Latimeria): Slow rate of evolution or recent
speciation? Marine Biology, 157(10), 2253–2262. https:// doi. org/ 10.
1007/ s00227‑ 010‑ 1492‑7
Toriño, P., Soto, M., & Perea, D. (2021). A comprehensive phylogenetic analy‑
sis of coelacanth fishes (Sarcopterygii, Actinistia) with comments on
the composition of the Mawsoniidae and Latimeriidae: Evaluating old
and new methodological challenges and constraints. Historical Biology,
33(12), 3423–3443. https:// doi. org/ 10. 1080/ 08912 963. 2020. 18679 82
von der Weid, J. (1960). Géologie des Préalpes médianes au SW du Moléson
Préalpes fribourgeoises. Eclogae Geologicae Helvetiae, 53(2), 523–624.
von Huene, F. (1939). Ein ganzes Ichthyosaurierskelett aus den westsch‑
weizerischen Voralpen. Mitteilungen der Naturfoschenden Gesellschaft in
Bern, 1939, 1–14.
Weidmann, M. (1981). Un lchtyosaure dans le Lias supérieur des Préalpes
médianes vaudoises. Bulletin de la Société Vaudoise des Sciences
naturelles No. 359, Vol. 75. Fasc., 3, 165–170.
Weidmann, M. (1993). Feuille 1244 Châtel St. Denis. Atlas géol. Suisse 1:25
000. Notice Explicative, 92, 1–57.
15 Page 20 of 20
C.Ferrante et al.
Witzmann, F., Dorka, M., & Korn, D. (2010). A juvenile Early Carboniferous
(Viséan) coelacanth from Rösenbeck (Rhenish Mountains, Germany)
with derived postcranial characters. Fossil Record, 13, 309–316. https://
doi. org/ 10. 1002/ mmng. 20100 0004
Woodward, A. S. (1890). Notes on name ganoid fishes from the English
Lower Lias. Annals and Magazine of Natural History, London, Series, 6(5),
430–436.
Woodward, A. S. (1891). Catalogue of Fossil Fishes in the British Museum (Natu-
ral History). London: Printed by order of the Trustees.
Woodward, A. S. (1909). The Fossil Fishes of the English Chalk. Part 5. London:
London Palaeontographical Society.
Publisher’s Note
Springer Nature remains neutral with regard to jurisdictional claims in pub‑
lished maps and institutional affiliations.
