Content uploaded by Saverio Bartolini-Lucenti
Author content
All content in this area was uploaded by Saverio Bartolini-Lucenti on Nov 27, 2018
Content may be subject to copyright.
DST
f OSSILIA- PORTS IN PAEONTOLOGY
Online publication of the Dept. of Earth Science of the University of Florence
sin,
Deinogalerix
ISBN 979-12-200-3408-1
9 791220 034081
Fossilia, Volume 2018: 1-2
Special SeSSion on Science communication
Short note
Istituto Nazionale di Fisica Nucleare, Sezione di Genova, Via Dodecaneso 33, 16146 Genova, Italy
Science and society have a stronger link than one can
see at a first glance, and we do not mean “applied scien-
ces”, whatever this can mean, but also “fundamental
sciences”. This can be explained in both directions.
Scientists get a significant part of their financing from
public institutions, States, local administrations, EU et
cetera, on one side, and this is is quite evident: this
also trigger a very common question, i.e. the infamous
“what is this thing useful for?”. On the other side, so-
ciety achieve a continuous improvement in the qua-
lity of life for citizens thanks to the advancement in
technology boosted by scientific research. This link is
perceived as looser than the other, and this is related to
the poor communication from the scientific commu-
nities to the citizens: the importance of proper scien-
tific communication for the general public has been
recognised in several context and the effort to fill this
gap is gaining momentum with increasing speed in the
very last years. The parallel increase in importance of
internet is opening new channels for scientific dissemi-
nation, parallel to the traditional ones. We all grew up
(at least til my generation) watching TV and reading
books or specialised press, when we wanted to approa-
ch science as “non experts”: both these activities were
mainly performed by communication professionals
with a strong science background and the support of
scientists. Nowadays the challenge is to have the lar-
gest number of scientists involved in scientific commu-
nication, ready to give an interview, write a dissemina-
tion paper, produce a “content”, which can be audio,
video, interactive and so on.
The focus, therefore, is moving from the communi-
cation professional to the scientist. What a scientist
can do to play a succesful role in this game? I have
gained some experience in the field thanks to Istituto
Nazionale di Fisica Nucleare, Pint of Science and spe-
cially Scientificast, this experience can be summarised
in few points.
First of all, science is of interest for many unsu-
spectable people. Trying to make it accessible is defi-
nitely worth of. Many people will thank you for your
efforts.
Never presume that something will be of no interest
for the public. Scientificast had a broad success for blog
posts concerning fundamental physics, number theory
and the strangest topics. Also the usual consideration,
“it is too complicated to be explained in a reasonable
way for the general public should be avoided”.
This is the most critical aspect. A scientist is used to
some jargon that is hardly comprehensible (when not
“wrong”) to the public. Often he jumps into the core
of the matter skipping a proper introduction, assuming
that everyone has a knowledge background somehow
comparable to the one he has. This is usually false and
also the idea of being able to speak to anyone is false:
Science and society: let’s stay in touch
Andrea Bersani
Bullet-pointS aBStract
• Science and society have a very strong link and we, as scientists, should consi-
der how to improve our communication and dissemination skills to implement
a positive feedback;
• New internet-based media provide an unprecedented opportunity to be give a
significant contribution for all the scientists never involved in traditional com-
munication;
• The public is variegated and unpredictable, but interested and curious: we can
reach (and satisfy) a large audience with a relatively small effort
• The experience of Scientificast, Italian blog and podcast, is a successful exam-
ple of new media exploitation for science communication, with thousands of
people involved for each post or episode;
• We identified as keys for our success the involvement of young, motivated
scientists with a well-defined target and style, a format that can be replicated,
modified and improved by other groups.
KeywordS:
Science Communication;
Podcast;
Blog;
New Media.
How to cite: Bersani (2018). Science and society: let’s stay in touch. Fossilia, Volume 2018: 1-2. https://doi.org/10.32774/
FosRepPal.20.1810.S10102
Fossilia - Reports in Palaeontology
Invited Paper - Short Note: Bersani
2
a scientist should always have in mind a target when
he decides to communicate science. Usually a good
compromise is thinking of people with a high scho-
ol diploma, but this is not the only possible choice.
There is an increasing interest in science communica-
tion and scientists’ role in this activity is getting more
and more relevant: any scientist should consider to de-
vote part of his time in science dissemination, exploi-
ting all the communication media available nowadays.
Andrea Bersani is from Genova, where he works as tech-
nology scientist at Istituto Nazionale di Fisica Nucleare.
Since 2013 he collaborates with Scientificast, one of the
most relevant Italian scientific podcasts in Italy, both
as a podcaster and as a blogger. He is a co-founder of
Pint of Science Italia and has collaborated with Festival
della Scienza, FameLab and other science dissemination
initiatives.
Fossilia, Volume 2018: 3-5
Master in Science Communication - Interdisciplinary Laboratory For Advanced Studies (ILAS) - International School for Advanced Studies
(SISSA) via Bonomea 265, 34136 Trieste TS; atavecch@sissa.it
Special SeSSion on Science communication
Short note
Science is - in principle - the ultimate source of
reliable information. No other institutions in mo-
dern society can be relied upon to produce credible
knowledge, especially when issues are politicized. If
science communication is tainted by special interests,
if it is confused with persuasion, if there is constant
suspicion of bias, this harms not only informed deci-
sion-making, but also the institution of science itself.
Science communication, in the most general sense, is
the crucial connection between the world of science
production and the rest of society. Therefore, the cre-
dibility of science as an institution is itself dependent
on the credibility of science communication.
Science communication is an evolving ecosystem,
with many stakeholders contributing to the creation
of shared meaning about scientists and science. To un-
derstand the recent growth of science communication
endeavours must also understand the goals and inte-
rests involved. More and more frequently financing
institutions explicitly require science communication
efforts to be included in grant proposals. For instance,
the follow up to the Horizon 2020 European research
framework - the 100 billion euros Horizon Europe ini-
tiative - requires all grant proposals to include a work
package on both Public Engagement/Dissemination
(PE) and Responsible Research and Innovation (RRI)
efforts to receive funding. It is one of the many ways in
which scientists are under a growing pressure to com-
municate: some observers have noted that the already
broken “publish or perish” system is changing towards
an even more demanding “publish and be social or pe-
rish” paradigm. Nevertheless, most institutions do not
provide any specific training in science communica-
tion for their scientists.
While, in theory, it is reasonable to expect scientist
to be the best communicators of their own research
work, in practice this expectation often turns out to
be false, due to the competition for public attention.
Scientists involved in science communication can
sometime reach wide popularity, very often indepen-
dently from the value of their research, and more often
than not as controversial figures. Inside the scientific
community there is still a strong ambivalence towards
those who receive media attention due to science com-
munication efforts. This phenomenon, the so-called
“Gould Effect” (named for the paleontologist Stephen
Jay Gould), is becoming ever more important due to
impact of social media and the rise of quantitative
indicators of reputation like altmetrics. More impor-
tantly, from a science communication perspective, the-
se “visible scientists” are the main actors in the con-
struction of shared narratives about what it means to
be a scientist and, consequently, about the nature of
science itself.
The idea that scientists ought to communicate with
the largest possible public has quickly reached enor-
mous popularity, due to being aligned both with ap-
peals for a more open and reliable science and, at the
same time, with the needs of political legitimation and
institutional PR. The problem of justifying scientific
funding has been deferred to universities and research
Bullet-pointS aBStract
• The credibility of science as an institution is itself dependent on the credibility
of science communication.
• Science communication is best understood as an ecosystem, with many sta-
keholders and forces interacting and contributing to the construction of shared
meanings about science and scientists
• The current science communication ecosystem is characterized by the loss of
trusted gatekeepers and by the blurring between different genres of communi-
cation (especially self-promotion, advertising and informational content).
• There is very little to no correlation between lay public knowledge about scien-
ce and attitudes towards science; trust is very often the critical factor.
KeywordS:
Science popularization;
society;
credibility;
communication ecosystem.
The science communication ecosystem
Alessandro Tavecchio
How to cite: Tavecchio (2018). The science communication ecosystem. Fossilia, Volume 2018: 3-5. https://doi.org/10.32774/
FosRepPal.20.1810.S20305
Fossilia - Reports in Palaeontology
4
centers that have to document the quality of their re-
search and their spending as “public accountability”.
Outreach and competition for attention are more often
than not already understood to be a central element of
academic activity and inherently desirable.
The competition for attention from a mainstream
audience means institutions often rely on a “push
communication mode”, very different from the parti-
cipatory and democratic rhetoric used to justify the
importance of science communication. Inevitably, this
kind of practice mixes institutional propaganda with
informative comment, being economically motivated
by branding, image building and marketing. Conse-
quently, since the early 2000s, new stakeholders have
risen in the ecosystem of science communications:
press officers and public relation specialists, involved
in corporate communication for Universities and re-
search centers.
Another fundamental stakeholder in the science
communication ecosystem are science journalists.
Since the early 20th century, science journalism has
gone through deep changes in philosophy and in bu-
siness practices, transforming science journalists from
“translators” of frontier research, to promoters of
science literacy, to critical observers and commenters.
Now more than ever the role of science journalists is
changing for two related reasons: the death of tradi-
tional print journalism business models and the ascent
of social media have stripped journalists of their “tru-
sted middleman” or “gatekeeper” role, creating new
direct avenues of communication between knowled-
ge production and society. While, in theory, science
journalists should be the most disinterested actor (in
the sense of independence from special interests) and
consequently a reliable source of science information,
or even a supervisor and “watchdogs” over the practi-
ces of the scientific community, economic and social
pressures have brought on the rise of “churnalism”
(the uncritical spreading of PR material), sensationa-
lization, clickbait, personalization and political bias in
scientific journalism. While scientists in general con-
tinue to be among the most trusted professionals in
surveys, trust in journalists has collapsed in the last 40
years, and now they are at the bottom of the rankings
together with politicians and bankers.
Unfortunately, some studies seem to indicate that the
trust in scientist and science is decreasing, and, for the
first time since the sixties, the trend is downward. This
is particularly visible when looking at privately funded
and industry funded scientists, but the pattern holds
true even for publicly-funded research.
Generally speaking, the contemporary science com-
munication ecosystem is characterized by the loss of
trusted gatekeepers and by the blurring between dif-
ferent genres of communication (especially self- pro-
motion, advertising and informational content). The
credibility of communication and the trust in the com-
municator are the main instruments of persuasion,
and their importance in science communication ef-
forts cannot be understated. Navigating this complex
ecosystem and communicating science effectively re-
quires well defined goals and careful planning.
Fig. 1. While trust in scientist is still high compared to most other professions and categories, Europeans feel that scientists
cannot be trusted to tell the truth about controversial scientific and technological issues because they depend more and more on
money from industry. This very specific erosion of trust is more dangerous than lack of knowledge, and the driving force behind
most recent changes in the science communication ecosystem. From the Special Eurobarometer 340 On Science And Technolo-
gy – European Commission (2015).
Invited Paper - Short Note: Tavecchio
The science communication ecosystem 5
referenceS
Guenther L. & Weingart P. (2016). Science commu-
nication and the issue of trust. Journal of Science
Communication, 15 (05): C3.
Marcinkowski F. & Kohring M. (2014). The changing
rationale of science communication: a challenge
to scientific autonomy. Journal of Science Commu-
nication, 13 (3): 1–8.
Peters H. P. (2013). ‘Gap between science and media
revisited: Scientists as public communicators’.
PNAS, 110 (Supplement 3): 14102–14109.
Shipman M. (2014). ‘Public relations as science com-
munication’ - Journal of Science Communication, 13
(03): C05.
Alessandro Tavecchio is a science communicator. As a freelan-
ce science journalist he writes for Motherboard, Vice, Wired and
others on issues spanning from biotechnology to open access to
science frauds. He’s also a founding member of the science popula-
rization initiative “Italia Unita per la Scienza”, blogger, podcaster,
social media manager for Science festivals, press officer and content
developer for Fondazione Telethon. He’s also completing a post-
graduate masters in Science Communication at the International
School For Superior and Advanced Studies in Trieste.
Fossilia, Volume 2018: 7-10
How to cite: Bartolini Lucenti (2018). Nyctereutes Temminck, 1838 (Mammalia, Canidae): a revision of the genus
across the Old World during Plio-Pleistocene times. Fossilia, Volume 2018: 7-10. https://doi.org/10.32774/FosRep-
Pal.20.1810.010710
Fossilia - Reports in Palaeontology
Nyctereutes
Temminck, 1838 (Mammalia, Canidae): a
revision of the genus across the Old World
during Plio-Pleistocene times
Saverio Bartolini Lucenti
Dottorato di Ricerca in Scienze della Terra, Università di Pisa, Via S. Maria 53, 56126 Pisa, Italy; saverio.bartolini@dst.unipi.it
Dipartimento di Scienze della Terra, Università degli Studi di Firenze 50121, Italy.
introduction
Nyctereutes Temminck, 1838 is nowadays represen-
ted by the single species Nyctereutes procyonoides (Gray,
1834). This taxon inhabits two separated areas of
Eurasia (Ward & Wurster-Hill, 1990): a natural one
(spanning from the Eastern part of Russia, China, Ko-
rean peninsula and Japan) and an artificial one resul-
ted from the accidental or intentional introduction du-
ring the 1920s and 1930s across Eastern and Central
Europe (from Finland to Hungary and from France to
the Caucasus). On the contrary, the fossil record reve-
als a much greater diversity as up to 10 species have
been described in literature (Bates, 1937; Rook et al.,
2017). Furthermore, the fossil distribution range was
not limited to Eurasia since Nyctereutes spp. were reco-
vered also from several regions of Africa (see among
others Werdelin & Dehghani, 2011).
The dentognatic features possessed by N. procyonoi-
des are diagnostic for the genus and, at least part of
them, were acquired by the extant raccoon-dogs’ ance-
stors as the result of progressive adaptations towards
a hypocarnivorous diet. Among these characteristics,
there are the reduction of slicing portions of the car-
nassials and enlargement of the crushing surface of
the molars, the developed subangular lobe, the expan-
sion of the angular process, and the development of
the insertion areas of the muscle pterigoideus media-
lis, on the medial side of the mandible.
Thanks to a considerably large sample from more
than twenty-five sites across the Old World, this stu-
dy combines the summary of the state of art of the
past research on the genus Nyctereutes together with the
most recent findings in the Old World.
materialS and methodS
The fossil sample studied for this research comes
from numerous Eurasian and African localities, hou-
sed in several institutions (See Table 1). The conside-
red species are Nyctereutes abdeslami Geraads, 1997,
?Nyctereutes barryi Werdelin & Dehghani, 2011, Nycte-
reutes donnezani (Depéret, 1890), Nyctereutes megama-
stoides (Pomel, 1842), Nyctereutes sinensis (Schlosser,
1903), Nyctereutes tingi Tedford & Qiu, 1991, Nyctereu-
tes terblanchei Ficcarelli et al., 1984, Nyctereutes vulpinus
Soria and Aguirre, 1976. The extant comparative sam-
ple includes specimens of N. procyonoides, Vulpes vulpes
(Linnaeus, 1758), Vulpes lagopus (Linnaeus, 1758) and
Cerdocyon thous Smith, 1839 housed in the MZUF and
AMNH (See abbreviation below).
Institutional abbreviations
AMNH, American Museum of Natural History, New
York (U. S. A.); AUT, Aristotle University Thessalo-
niki (Greece); GNM, Georgian National Museum,
Tbilisi (Georgia); IGF, Geological and Paleontologi-
cal Section of the Natural History Museum of the
University of Florence; MdC, Musée des Confluen-
ces, Lyon (France); MNCN, Museo Nacional de
Ciencias Naturales, Madrid (Spain); MNHN, Musée
National d’Histoire Naturelle, Paris (France); MZUF,
La Specola, Zoological section of the Natural History
Museum of the University of Florence; UCBL-1,Uni-
versité Claude-Bernard Lyon-1, Lyon (France).
diScuSSionS
The Old world record of Nyctereutes: an update
The earliest occurrences of the genus is in the Yushe
Bullet-pointS aBStract
• Although Nyctereutes is nowadays monospecific, the fossil record suggests a
larger diversity and wider distribution.
• The study of large sample from numerous sites of the Old World allowed a
critical revision at an intercontinental scale.
• Some specimens of Layna and the sample from Çalta reveal peculiar morpho-
logies, contrasting with the attribution to N. donnezani.
• The peculiarities of two samples testify to the urge for new research on fossil
Nyctereutes.
KeywordS:
Raccoon-dogs;
State of art;
Pliocene;
Biogeography;
Canidae.
Bartolini Lucenti
8
basin (China, ~MN14-15, Tedford et al., 2013) with
the primitive species N. tingi (Fig. 1A). Its morpholo-
gies strongly differ from all the other Nyctereutes spp.
for the modest or scarce development of mandibular
and dental features (as pointed out by Tedford & Qiu,
1991).Around 4-3.5 Ma, there is a first burst in diver-
sity (Fig. 1B): i) firstly, the African record starts at this
time with the dubious and primitive ?N. barryi from
the Upper Laetoli Bed (Tanzania); ii) secondly, the oc-
currence of N. sinensis, in the same Chinese basin of
N. tingi, which marks the appearance of a taxon that
resembles the extant N. procyonoides for the degree of
developmente of its dentognatic features. Then in We-
stern Europe, N. donnezani was described from French
and Spanish sites. Some authors deem that this first
European species corresponds to N. tingi (see Tedford
& Qiu, 1991). In contrast to the Asian record (see Te-
dfor et al., 2013), during the late Pliocene in Europe,
primitive and derived forms of raccoon-dogs never co-
existed: N. donnezani was replaced by N. megamastoides.
A second burst is dated between 3.5-2 Ma (Fig. 1C).
N. megamastoides appears in Europe at this time (Barto-
lini Lucenti, 2017), and, partially due to its large range
of distribution (from Spain to Georgia, Rook et al.,
2017), it is one the most renowned fossil species of
this genus. The retention of strongly derived dental
and cranio-mandibular features in N. megamastoides,
led some scholars to suggest the idea that this taxon
and N. sinensis are closely related (Soria & Aguirre,
1976), although some morphological differences re-
main (Rook et al., 2017). The sample from St. Vallier
(France), historically attributed to N. megamastoides
(Viret, 1954; Martin, 1971) should be classified as a se-
parate taxon from N. megamastoides and in this respect,
for the presence of some features resembling the genus
Vulpes, (Soria & Aguirre, 1976) suggested for this re-
mains the attribution to the species N. vulpinus. Apart
from the controversial taxon ?N. barryi , the African
record of Nyctereutes is relatively scanty but diverse.
Younger although still early Pliocene in age, the pri-
mitive Nyctereutes lockwoodi Geraads et al., 2010 from
Dikika (Ethiopia) shows several peculiar morpholo-
gies, which justify the separation in a different species.
Other findings in Africa are the Early Pleistocene N.
abdeslami from Morocco, and the South African N. ter-
blanchei (both derived species).
After 2 Ma, and during the last part of the Early Plei-
stocene, the findings of Nyctereutes across the whole
Old World become more and more scarce and the di-
versity of the genus declined. The first record of the
extant N. procyonoides appears in Middle Pleistocene
deposits of China (Zhoukoudian 1 and 13, Tedford &
Qiu, 1991) and remained the only species of the ge-
nus until today, with the exception of the Palestinian
Nyctereutes vinetorum Bates, 1937 (which may simply
represent a larger form of the extant species).
The record of Layna and Çalta
The ongoing revision of the material from two im-
portant localities referred to MN15 (Layna, Spain;
Çalta, Turkey) yielded some unexpected results. In li-
terature, these samples are referred to N. donnezani (So-
ria & Aguirre, 1976; Ginsburg, 1998). Nevertheless, at
Layna few specimens show morphological similarities
and morphometric proportions that cannot be explai-
ned as intraspecific variability, as they suggests affinity
with derived forms. The Nyctereutes from Çalta posses-
ses an undeniable primitive-like morphology of the
mandible corpus with a reduced subangular lobe (as
noted by Ginsburg, 1998) albeit its dental proportions
or the shape of the molars (especially upper ones) does
not fit with the attribution to N. donnezani. On the con-
trary, the dental characters showed by the Çalta Nycte-
reutes are close to those of forms like N. sinensis, N. vul-
pinus or, even more, N. megamastoides.
concluSionS
The fossil record of the genus Nyctereutes, spanning
in whole Old World in the last 5 Ma, testifies to a great
variety in morphologies and adaptations in diet. Many
scholars have investigated the evolutionary history of
this genus, yet many issues are still matters of debate
and fertile grounds for research (e.g., the phylogenetic
relationships between the species). This short over-
Species Locality Repository
N. tingi Yushe Basin (Liujiagou,
Nanzhuanggou) (China);
Megalo Emvolon (Greece)
AMNH
N. sinensis Yushe Basin (Beihai,
Liujiagou, Xiachuang,
Zhangwagou, Zha-
ozhuan), Nihewan Basin
(China)
AMNH,
MNHN
N. donnezani Perpignan-Roussillon
(France); Layna, La Gloria
4* (Spain)
AMNH,
MdC, MNCN,
UCBL-1
N. vulpinus St. Vallier (France) Mdc, UCBL-1
N. megamastoides Senéze, Perrier-L'Etouaires
(France); Kvabebi (Geor-
gia); Dafnero (Greece);
Csarnota, Beremend
(Hungary); Montopoli, S.
Giusto (Italy); El Rincon,
Villarroya (Spain)
AUT, GNM,
HMNH,
IGF, MNCN,
MNHN,
UCBL-1
N. abdeslami Ahl al Oughlam
(Morocco)
MNHN
N. terblanchei Kroomdrai A*
(South Africa)
IGF
?N. barryi Upper Laetoli Bed*
(Tanzania)
MNHN
Nyctereutes sp. Çalta (Turkey) MNHN
Tab. 1. List of the considered sample for this study: the
species of Nyctereutes, the localities where they come from
and the institutions in which they are housed in.
Nyctereutes
across the Old World during the PLio-PLeistocene 9
view of Nyctereutes spp. summarizes our current know-
ledge of their diversity and evolutionary history, and
points out peculiar two cases (i.e. those of Layna and
Çalta), which certainly require deep studies in the fu-
ture. Should the hypothesis of an early appearance of
form with mixed pattern of morphologies in the MN15
be validated, it could reveal the presence in the Ear-
ly Pliocene of an alternative lineage of raccoon-dogs,
with serious implications on Nyctereutes dispersion and
evolution across the Palearctic, the Afrotropical and
the Indomalayan regions.
acKnowledgementS
The author is thankful to the kindness and availability of E.
Cioppi (IGF), P. Agnelli (MZUF), M. Bukhsianidze (GNM),
E. Robert (UCBL-1), D. Berthet (MdC), J. Galkin and J.
Meng (AMNH), M. Gasparik (HMNH), C. Argot (MNHN),
S. Fraile (MNCN), and G. Koufos (AUT). This research has
been partly supported by the SYNTHESYS Project http://
www.synthesys.info/ (Project Numbers ES-TAF-6553, HU-
TAF-6520), which is financed by European Community
Research Infrastructure Action under the FP7 “Capacities”
Program.
referenceS
Bartolini Lucenti S. (2017). Nyctereutes megamastoides (Cani-
dae, Mammalia) from the early and middle Villafran-
chian (late Pliocene and early Pleistocene) of the Lower
Valdarno (Firenze and Pisa, Tuscany, Italy). Rivista Ita-
liana di Paleontologia e Stratigrafia, 123: 211-218.
Bates D. M. A. (1937). Palaeontology: The fossil fauna of
the Wady el-Mughara caves. In The Stone Age of Mount
Carmel. Excavations at the Wady El-Mughara. Oxford
(Clarendon Press), vol. 1, part 2: 137-233.
Depéret C. (1890). Les Animaux Pliocènes du Roussillon.
Mémoires de la Societé Géologique de France, 3: 5–195.
Ficcarelli G., Torre D. & Turner A. (1984). First evidence for
a species of raccoon dog, Nyctereutes Temminck, 1838, in
South African Plio-Pleistocene deposits. Boll. Soc. Pale-
ont. It., 23: 125-130.
Geraads D. (1997). Carnivores du Pliocène terminal de Ahl
al Oughlam (Casablanca, Maroc). Géobios, 30: 127- 164.
Geraads D., Alemseged Z., Bobe R. & Reed D. (2010). Nycte-
reutes lockwoodi, n. sp., a new canid (Carnivora: Mamma-
lia) from the middle Pliocene of Dikika, Lower Awash,
Ethiopia. Journal of Vertebrate Paleontology, 30: 981-987.
Ginsburg L. (1998). Le gisement de vertébrés pliocènes de
Çalta, Ankara, Turquie. 5. Carnivores. Geodiversitas, 20,
Fig. 1. Geographic and temporal distribution of Nyctereutes spp. from the earliest Early Pliocene to present times across the Old
World. A, earliest Early Pliocene, 5.3-4.5 Ma. B, Early-Late Pliocene, 4.5-3.5 Ma. C, Latest Pliocene-Early Pleistocene (partim),
3.5-1.5 Ma. D, Middle Pleistocene-Present times, 800-0 ka.
Bartolini Lucenti
10
379-396.
Gray J. (1834). Illustration of Indian zoology, consisting of
coloured plates of new or hitherto unfigured Indian ani-
mals from the collection of Major General Hardwicke.
Fol, London 2: pI. 1.
Linnaeus C. (1758) - Systema Naturae per regna tria natu-
rae, secundum Classes, Ordines, Genera, Species, cum
characteribus, differentiis, synonymis, locis. Tomus I,
10th edition: Holmiae, Laurentius Salvius, Stockholm,
Sweden, 824 p.
Martin R. (1971). Les affinités de Nyctereutes megamastoi-
des (Pomel), canidé du gisement villa franchien de
Saint-Vallier (Drôme, France). Palaeovertebrata, 4: 39-58.
Pomel M. (1842). Nouvelle espèce de chien fossile découv-
erte dans les alluvions volcaniques de l’Auvergne. Bulle-
tin de la Société Géologique de France,14: 38-41.
Rook L., Bartolini Lucenti S., Bukhsianidze M. & Lordiki-
panidze D. (2017). The Kvabebi Canidae record revisi-
ted (late Pliocene, Sighnaghi, eastern Georgia). Journal
of Paleontology, 91: 1258–1271.
Schlosser M. (1903). Die fossilen Säugethiere Chinas nebst
einer Odontographie de recenten Antilopen. Abhandlun-
gen der Mathematisch-Physikalischen Klasse der Königlich
Bayerischen Akademie der Wissenschaften, 22: 1-220.
Smith C. H. (1839). The Canine Family in general or the
genus Canis. In W. Jardine (ed), The naturalist’s library,
vol. 18. Natural history of dogs, vol. 1. Edinburgh: W.
H. Lizars, 267 pp.
Soria D. & Aguirre E. (1976). El cánido de Layna: revisión
de los Nyctereutes fósiles. Trabajos Neógeno y Cuaternario,
5: 83-115.
Tedford R. H. & Qiu Z. (1991). Pliocene Nyctereutes (Carni-
vora: Canidae) from Yushe, Shanxi, with comments on
Chinese fossil raccoon-dogs. Vertebrata PalAsiatica, 29:
176-189.
Tedford R. H., Qiu Z. X. & Flynn L. J., eds. (2013). Late Ce-
nozoic Yushe Basin, Shanxi Province, China: Geology
and Fossil Mammals. Springer, New York.
Temminck C. (1838). Over de kennis en de Verbreiding der
Zoogdieren van Japan. Tijdschrift Natuurlijke Geschie-
deins en Physiologie, 5: 273- 293.
Viret J. (1954). Le loess à bancs durcis de Saint-Vallier
(Drôme) et sa faune de mammifères villafranchiens:
Nouveau Archives du Musee d’Histoire Naturelle de Lyon, 4:
1–200.
Ward O. G. & Wurster-Hill D. H. (1990). Nyctereutes procyo-
noides. Mammalian Species, 358, 1-5.
Werdelin L. & Dehghani R. (2011). Carnivora. In Harrison
T. (ed.) Geology and Paleontology of Laetoli: Human
evolution in context. Springer, New York: 189-232.
Manuscript received 15 July 2018
Received after revision 14 September 2018
Accepted 20 September 2018
Fossilia, Volume 2018: 11-12
Preliminary report on the taxonomic revision of
Fossil Equidae from Tizi’n Tadderht (Ouarzazate,
Morocco)
Omar Cirilli1 & Samir Zouhri2
1 Dipartimento di Scienze della Terra, Università degli Studi di Firenze 50121, Italy; omar.cirilli@yahoo.com
2 Laboratoire de géosciences, faculté des Sciences, Université Hassan II-Casablanca, Morocco; samirzouhri@yahoo.fr
introduction
We are reviewing the sample of fossil Equidae, belon-
ging to the genus “Hipparion” s.s. (see Armour-Chelu
& Bernor, 2011) from Tizi’n Tadderht (Ouarzazate,
Morocco), a fossiliferous site chronologically referable
to the Late Miocene.
The fossiliferous locality of Tizi’n Tadderht, already
known in the literature (Geraads et al., 2012; Zouhri et
al., 2012), has yielded a small but significant vertebrate
fossil association. It represents the first documentation
of a Late Miocene vertebrate fauna in the western area
of North Africa. This new fauna allows to expand the
possibilities of investigation on the biogeographical
and evolutionary record of the vertebrate fossils in the
circum-Mediterranean area.
materialS and methodS
The group of fossil Equidae under revision has been
preliminarly studied by Zouhri et al. (2012), who
identified the following hipparionini species: aff. Cre-
mohipparion periafricanum (Villalta & Crusafont, 1957),
Hippotheriini gen. et sp. indet., and cf. Hippotherium
primigenium (Von Meyer, 1829).
The sample retrieved from the considered area is
under review through the description of the external
anatomy morphologies and the dimensional measu-
rements analysis. In addition, a comparison with the
collection of fossil Equidae of the Libyan site of As
Sahabi is also on the way. At As Sahabi the following
species of Equidae hipparionini are represented (Ber-
nor et al., 2008; 2012): Sivalhippus sp., Eurygnathohip-
pus feibeli Bernor & Harris, 2003 and Cremohipparion
matthewi (Abel, 1926).
diScuSSionS
This revision of the Tizi’n Tadderht association led
us to confirm the identification of three hipparionine
species, differing by size and anatomical details.
The largest form has been identified by Zouhri et al.
(2012) as belonging to Hippotherium sp.; specimens
referred to this taxon show some anatomical features
suggesting an attribution to the genus Hippotherium
Kaup, 1833, such as having all tooth elements isolated
and not included in the cementum; a well developed
mesostyle, complex plications of the pre- and postfos-
settes, an anterostyle more developed and isolated.
A medium-sized Equidae has been assigned to Hip-
potheriini gen. et sp. indet by Zouhri et al. (2012). The
revision of the entire collection attributable to this mi-
dlle-sized hipparionine horse is under way. We are te-
sting the hypothesis that such a form could be attribu-
ted to the genus Eurygnathohippus (Van Hoepen, 1930)
(Fig. 1). This latter taxon is present within the fossil
assemblage of As Sahabi (Libya) and Tizi’n Tadderht
specimens are remarkably similar in dimensions and
proportions to E. feibeli from the Libyan site.
Finally the few fragmentary fossils representing a very
small hipparionine horse from Tizi’n Tadderht have
been described by Zouhri et al. (2012) as Cremohip-
parion aff. C. periafricanum. Interestingly a very small
hipparionine species also occurs in the Late Miocene
site of As Sahabi (where it has been identified as Cre-
mohipparion matthewi by Bernor et al., 2008). Our com-
parisons confirm the attribution of this small sample
from Tizi’n Tadderht to the genus Cremohipparion Qiu
et al., 1987. A deeper comparison with the As Sahabi
sample is necessary before any conclusion for a deter-
mination at the specific level.
Bullet-pointS aBStract
• The fossiliferous locality of Tizi’n Tadderht is a significant latest Miocene
vertebrate fossil association in the North Africa.
• The fossil Equidae from Tizi’n Tadderht had been preliminarly studied (Zouhri
et al. 2012).
• We prelimiarly revise the Tizi’n Tadderht Equidae association in order to have
new insights on the genus Hipparion in Morocco.
KeywordS:
Hipparion;
Tizi’n Tadderth;
Sahabi;
Morocco;
Libya.
Corresponding author email: omar.cirilli@yahoo.com
How to cite: Cirilli & Zouhri (2018). Preliminary report on the taxonomic revision of Fossil Equidae from Tizi’n Tadderht
(Ouarzazate, Morocco). Fossilia, Volume 2018: 11-12. https://doi.org/10.32774/FosRepPal.20.1810.021112
Fossilia - Reports in Palaeontology
Cirilli & Zouhri
12
concluSionS
The re-evaluation of the hipparionine equids assem-
blage from Tizi’n Tadderht site is of particular impor-
tance as it extends the paleogeographic record of Hip-
parion and other hipparionene species present in other
African sites, showing that this clade is well represen-
ted in the African fossil record of the Late Miocene.
At the same time, an exhaustive comparative study of
the fossil Hipparionini assemblages from the As Saha-
bi site (Libya) and from Tizi’n Tadderht site (Moroc-
co), will allow us to contrast similar contemporaneous
faunal assemblages from the Late Miocene of North
Africa and to have a better understanding of the de-
velopment and the diffusion of Equidae assemblage
during the Late Miocene in North Africa (as well as
on the Late Miocene faunal exchange between Africa
and Europe).
referenceS
Abel O. (1926). Die Geschichte der Equiden auf dem Bo-
dem Noramerikas. Verhandlungen Zoologish-Botanischen
Gesellschaft, 74: 159-164
Armour-Chelu M. & Bernor R. L. (2011). Equidae. In Har-
rison T. (ed.), Paleontology and geology of Laetoli:
Human evolution in context (Fossil hominins and the
associated fauna, Vol. 2), Springer, Dordrecht: 295-326.
Bernor R. L. & Harris J. M. (2003). Systematics and evolu-
tionary biology of the Late Miocene and Early Pliocene
hipparionine horses from Lothagam, Kenya. In Har-
ris J.M. & Leakey M.G. (eds), Lothagam: The Dawn
of Humanity in Eastern Africa, Columbia University
Press, New York: 387-438.
Fig. 1. Hippotheriini gen. et sp. indet from Tizi’n Tadderht – MTE 12, right hemimandible, A, p2-m3 in occlusal view. B, hemi-
mandible in labial view. Scale bar 5 cm.
Bernor R. L., Kaiser T. M. & Wolf D. (2008). Revisiting
Sahabi equid species diversity, biogeographic patterns
and diet preferences. In El-Arnauti A., Salem M., Pa-
vlakis P. & Boaz N. (eds), Circum- Mediterranean Geo-
logy and Biotic Evolution During the Neogene Period:
The Perspective from Libya. Garyounis Scientific Bulletin,
Special Issue, 5: 159-167.
Bernor R. L., Boaz N. T. & Rook L. (2012) Eurygnathohip-
pus feibeli (perissodactyla: Mammalia) from the Late
Miocene of As Sahabi (Libya) and its Evolutionary and
Biogeographic Significance. Bollettino della Società Pale-
ontologica Italiana, 51 (1), 39-48.
Geraads D., El Boughabi S. & Zouhri S. (2012). A new ca-
prin bovid (Mammalia) from the late Miocene of Mo-
rocco. Paleontologia africana, 47:19-24.
Kaup J. J. (1833). Mitteilungen an Professor Bronn. Neues
Jahrbuch fur Mineralogie, Geognoise, Geologie und Petrefa-
ktenkunde Jahrgang, 1833: 419-420.
Qui Z. X., Huang W. L. & Guo Z. H. (1987). The Cinese
hipparionine fossils. Paleontologia Sinica, New Series 25:
1-150.
Van Hoepen E. C. N. (1930). Fossiele Pferde van Cornelia.
Paleontologiese Navorsing van die Nasionale Museum, Bloe-
mfontein, 2: 13-24.
Villalta, J. F. & Crusafont, M. (1957). Les gisements de
mammifères du Néogène espagnol. Comptes Rendus de la
Société Géologique de France, 9-10.
Von Meyer H.1829. Taschenbuch für die gesamte Mineralo-
gie. Zeitschrift Mineralien, 23: 150–152.
Zouhri S., Geraads D., El Boughabi S. & El Harfi A. (2012).
Discovery of an Upper Miocene Vertebrate fauna near
Tizi N’Tadderht, Skoura, Ouarzazate Basin (Central
High atlas, Morocco). Comptes Rendus Palevol, 11: 455–
461.
Manuscript received 30 July 2018
Received after revision 28 September 2018
Accepted 1 October 2018
Fossilia, Volume 2018: 13-14
A kogiid sperm whale from the lower Pliocene of
the Northern Apennines (Italy)
Alberto Collareta1 , Franco Cigala Fulgosi2 & Giovanni Bianucci1
1 Dipartimento di Scienze della Terra, Università di Pisa, via Santa Maria 53, 56126 Pisa, Italy; alberto.collareta@unipi.it
2 Strada Martinella 292, 43124 Parma, Italy
introduction
Among modern toothed whales (Cetacea: Odontoce-
ti), dwarf and pygmy sperm whales [Kogia sima (Owen,
1866) and Kogia breviceps (de Blainville, 1838)], re-
spectively) are the only living members of the physete-
roid family Kogiidae, known as diminutive and largely
unknown relatives of the great sperm whale (Physeter
macrocephalus Linnaeus, 1758). Extant kogiids inhabit
tropical to temperate open-sea environments outside
the Mediterranean Basin (e.g. McAlpine, 2002). The
fossil record of Kogiidae is to date represented by a
few skulls and more abundant isolated ear bones from
Neogene deposits of the Northern Hemisphere (e.g.
Barnes, 1973; Pilleri, 1987; Cigala Fulgosi, 1996; Bia-
nucci & Landini, 1999; Lambert, 2008; Whitmore &
Kaltenbach, 2008; Bianucci et al., 2011; Vélez-Juarbe
et al., 2015, 2016), with the significant exception of
the late Miocene record from the Pisco Formation
of southern Peru (de Muizon, 1988; Collareta et al.,
2017).
Here we report on a new fossil kogiid specimen col-
lected by one of us (F.C.F.) at S. Andrea Bagni (Parma
Province, Italy), a site where lower Pliocene marine
mudstones (“blue clays” sensu lato) are exposed. The-
se sediments have also yielded a rich deep-water ela-
smobranch assemblage [including teeth attributed to
rare squaloid sharks such as Scymnodalatias aff. garricki
Kukuev & Konovalenko, 1988, Scymnodon ringens Bar-
bosa du Bocage & Brito Capello, 1864, and Zameus
squamulosus (Günther, 1877)] which depict the presen-
ce of psychrospheric water masses of Atlantic origin
in the Adriatic palaeo-area during a phase of remar-
kable “oceanization” of the Mediterranean Basin (Ci-
gala Fulgosi, 1986; 1996).
materialS and methodS
The kogiid specimen from S. Andrea Bagni, which is
currently stored at Museo di Storia Naturale dell’Uni-
versità di Pisa as MSNUP I-17603, consists of a par-
tially complete cranium (lacking the basicranium and
the left part of the supracranial basin), one vertebra,
one fragment of rib, and one isolated tooth.
diScuSSionS and concluSionS
MSNUP I-17603 belongs to a new taxon of Kogii-
dae sharing various similarities with the recently de-
scribed Nanokogia isthmia Vélez-Juarbe et al., 2015,
from the upper Miocene of the Chagres Formation
of Panama. MSNUP I-17603 mainly differs from N.
isthmia by displaying two well-distinct fossae on the
right side of the supracranial basin (here interpreted
as accomodating the vocal chamber and the case for
the spermaceti organ, respectively) and a proportio-
nally longer rostrum; it further differs from the late
Pliocene Mediterranean species Kogia pusilla (Pilleri,
1987) by the larger size and a different architecture of
the supracranial basin. A low temporal fossa and the
absence of dental enamel may suggest that, like the
dwarf and pygmy sperm whales, MSNUP I-17603 was
a suction feeder rather than a raptorial predator; howe-
ver, the presence of an unusually long rostrum indi-
cates that the foraging technique of this extinct form
differed somewhat from that of modern kogiids, whi-
ch in turn feature the shorter rostrum among extant
odontocetes. Our preliminary phylogenetic analysis
recovers MSNUP I-17603 as a member of the subfa-
mily Kogiinae, which includes Kogia spp., Koristocetus
pescei Collareta et al., 2017, N. isthmia, and Praekogia
Bullet-pointS aBStract
• We report on a new specimen of Kogiidae from S. Andrea Bagni, a Zanclean
fossiliferous site of northern Italy.
• This specimen consists of a partially complete cranium, one vertebra, one
fragment of rib, and one tooth.
• The S. Andrea Bagni kogiid is recognized as representative of a new taxon in
the subfamily Kogiinae.
• Association of this specimen with teeth of deep-water squaloids provides inte-
resting palaeoecological hints.
KeywordS:
Adriatic palaeo-area;
Mediterranean Basin;
Physeteroidea;
palaeoecology;
psychrosphere.
Corresponding author email: alberto.collareta@unipi.it
How to cite: Collareta et al. (2018). A kogiid sperm whale from the lower Pliocene of the Northern Apennines (Italy).
Fossilia, Volume 2018: 13-14. https://doi.org/10.32774/FosRepPal.20.1810.031314
Fossilia - Reports in Palaeontology
COllareta et al.
14
cedronensis Barnes, 1973. Although Koristocetus is re-
covered as the earliest branching lineage of Kogiinae,
the relationships among Kogia Gray, 1846, Nanokogia
Vélez-Juarbe et al., 2015, Praekogia Barnes, 1973, and
MSNUP I-17603 are unresolved in our strict consen-
sus tree.
Since modern kogiids do not inhabit the Mediter-
ranean Sea, and considering also that they are belie-
ved to forage on deep-water cephalopods and fish at
or beyond the edge of the continental shelf, the as-
sociation of MSNUP I-17603 with a psychrospheric
elasmobranch assemblage could prove noteworthy.
Indeed, at the beginning of the Pliocene, the Gibraltar
connection was likely deep and controlled by estuari-
ne dynamics, thus allowing the passage of deep-water
organisms (some of which constitute the core of the
diet of extant Kogia) from the northern Atlantic Oce-
an to the Mediterranean Basin (Cigala Fulgosi, 1996,
and references therein). More generally, the finding of
MSNUP I-17603 suggests that our knowledge of the
evolutionary path of kogiid sperm whales is still far
from being exhaustive, as well as our understanding of
their past diversity, disparity, and distribution.
acKnowledgementS
Our gratitude to Chiara Sorbini, Felix G. Marx, Simone Ca-
sati, Aldo Marcelo Benites Palomino, and two anonymous
reviewers.
referenceS
Barnes L. G. (1973). Praekogia cedrosensis, a new genus and
species of fossil pygmy sperm whale from Isla Cedros,
Baja California, Mexico. Contributions in Science of the
Natural History Museum of Los Angeles County, 247: 1-20.
Bianucci G., Gatt M., Catanzariti R., Sorbi S., Bonavia C.
G., Curmi R. & Varola A. (2011). Systematics, biostra-
tigraphy and evolutionary pattern of the Oligo-Miocene
marine mammals from the Maltese Islands. Geobios, 44:
549-585.
Bianucci G. & Landini W. (1999). Kogia pusilla from the
Middle Pliocene of Tuscany (Italy) and a phylogenetic
analysis of the family Kogiidae (Odontoceti, Cetacea).
Rivista Italiana di Paleontologia e Stratigrafia, 105: 445-
453.
de Blainville H. M. D. (1838). Sur les cachalots. Annales
Françaises et Étrangères d’Anatomie et de Physiologie, 2:
335-337.
Barbosa du Bocage J. V. & Brito Capello F. (1864). Sur quel-
ques espèces inédites de Squalidae de la tribu Acanthia-
na Gray, qui fréquentent les côtes du Portugal. Procee-
dings of the Zoological Society of London, 1864: 260-263
Cigala Fulgosi F. (1986). A Deep Water Elasmobranch
Fauna from a Lower Pliocene Outcropping (Northern
Italy). In Uyeno T., Arai R., Taniuchi T. & Matsuura K.
(eds.), Indo-Pacific Fish Biology. Proceedings of the Se-
cond International Conference on Indo-Pacific Fishes,
Ichthyological Society of Japan: 133-139.
Cigala Fulgosi F. (1996). Rare oceanic deep water squaloid
sharks from the lower Pliocene of the Northern Apen-
nines (Parma Province, Italy). Bollettino della Società Pa-
leontologica Italiana, 34: 301-322.
Collareta A., Lambert O., de Muizon C., Urbina M. & Bia-
nucci G. (2017). Koristocetus pescei gen. et sp. nov., a di-
minutive sperm whale (Cetacea: Odontoceti: Kogiidae)
from the late Miocene of Peru. Fossil Record, 20: 259-
278.
Gray J. E. (1846). On the British Cetacea. Annals and Maga-
zine of Natural History, Series 1, 17: 82-85.
Günther A. (1877). Preliminary notes on new fishes col-
lected in Japan during the expedition of H.M.S. `Chal-
lenger’. Annals and Magazine of Natural History, Series 4,
20 (119): 433-446.
Kukuev E. I. & Konovalenko I. I. (1988). Two new species
of sharks of the genus Scymnodalatias (Dalatiidae) from
the North Atlantic and southeastern Pacific oceans. Vo-
prosy Ikhtiologii, 28 (2): 315-319. [In Russian]
Lambert O. (2008). Sperm whales from the Miocene of the
North Sea: a re-appraisal. Bulletin de l’Institut Royal des
Sciences Naturelles de Belgique, Sciences de la Terre, 78: 277-
316.
Linnaeus C. (1758). Systema Naturae sive Regna Tria Natu-
rae, secundum Classes, Ordines, Genera, Species, cum
characteribus, differentiis, synonymis, locis, Tomus I.
Editio decima, reformata. Laurentii Salvii, Stockholm,
824 pp.
McAlpine D. (2002). Pygmy and dwarf sperm whales. In
Perrin W.F., Würsig B. & Thewissen J.G.M. (eds.), En-
cyclopedia of marine mammals, Academic Press: 1007-
1009.
de Muizon C. (1988). Les Vertébrés de la Formation Pisco
(Pérou). Troisième partie: Les Odontocètes (Cetacea,
Mammalia) du Miocène. Travaux de l’Institut Français
d’Études Andines, 42: 1-244.
Owen R. (1866). On some Indian Cetacea collected by Wal-
ter Elliot, Esq. Transactions of the Zoological Society of
London, 6: 17-47.
Pilleri G. (1987). The Cetacea of the Italian Pliocene with a
descriptive catalogue of the species in the Florence Mu-
seum of Paleontology. Brain Anatomy Institute, Bern.
Vélez-Juarbe J., Wood A. R., De Gracia C. & Hendy A. J.
(2015). Evolutionary patterns among living and fossil
kogiid sperm whales: evidence from the Neogene of
Central America. PLOS ONE, 10: e0123909.
Vélez-Juarbe J., Wood A. R. & Pimiento C. (2016). Pygmy
sperm whales (Odontoceti, Kogiidae) from the Pliocene
of Florida and North Carolina. Journal of Vertebrate Pa-
leontology, 36: e1135806.
Whitmore Jr F. C. & Kaltenbach J. A. (2008). Neogene Ce-
tacea of the Lee Creek Phosphate Mine, North Caro-
lina. Virginia Museum of Natural History Special Publica-
tion, 14: 181-269.
Manuscript received 8 July 2018
Received after revision 4 October 2018
Accepted 5 October 2018
Fossilia, Volume 2018: 15-17
Stratigraphic paleobiology of an evolutionary
radiation: taphonomy and facies distribution of
cetaceans in the last 23 million years
Stefano Dominici1, Simone Cau2 & Alessandro Freschi2
1 Museo di Storia Naturale, Università degli Studi di Firenze, Firenze, Italy; stefano.dominici@unifi.it
2 Dipartimento di Scienze Chimiche, della Vita e della Sostenibilità Ambientale, Università degli Studi di Parma, Parma, Italy; cau.simo-
ne2@gmail.com, freschiales@gmail.com
introduction
The study of the stratigraphy and taphonomy of Neo-
gene cetaceans is a fundamental step to properly frame
the evolutionary radiation of this megafauna, at the
top of the pelagic marine ecosystem. Major evolutio-
nary steps have been summarised in recent studies, ta-
king the cetacean fossil record at face value and, whi-
le the emergence of modern adaptations during the
Neogene is not questioned (Berta et al., 2015; Marx
et al., 2016), still the comparison of diversity betwe-
en time intervals may be hampered by the quality of
the record, some evidence suggesting that some facies
are associated with a better fossil record than others
(Dominici et al., 2018). Nineteen cetacean families
are known in the Miocene, an epoch characterized by
the radiation of odontoceti and crown baleen whales.
During the Pliocene many large and small odonto-
ceti went extinct, while delphinids radiated, together
with balaenoidean and thalassotherian mysticeti,
two groups comprising today the largest tetrapods in
the history of life (Steeman et al., 2009; Marx et al.,
2015). The present study aims at collecting evidence
as to the facies distribution of the fossils upon which
these diversities have been calculated.
Changes in marine megafauna (MM) biodiversity ap-
pear to be strongly correlated with climate change.
An important extinction event has been identified in
the late Pliocene (between 3.8-2.6 Ma) and related to
enhanced climatic variability, higher-amplitude sea-le-
vel oscillations and loss of productive coastal habita-
ts (Pimiento et al., 2017). Cetaceans are the largest
animals among MM and form the largest part of the
MM fossil record (91 genus out of 215: Pimiento et
al., 2017). The foothills of the Northern Apennines
offer a particularly rich cetacean fossil record and an
area where available studies allow to explore this key
time of cetacean evolution at a stratigraphic resolution
finer than the stage. An increase in cetaceans diversity
is recorded around 3.2 – 3.0 Ma, in coincidence of
the mid-Piacenzian climatic optimum, and a drastic
decline at the Piacenzian-Gelasian boundary (Domi-
nici et al., 2018; Freschi et al., in press). Starting from
this evidence, the present study aims at exploring if
measured taxonomic diversity can be taken at face va-
lue, or if it is an artifact of the fossil record, by adding
data from the literature of Neogene cetaceans of other
geographic areas.
materialS and methodS
To check for a facies influence in the taphonomy of
Neogene cetaceans, we carried out a survey of 255 pa-
pers dealing with Neogene cetaceans reaching a global
picture to the exclusion of Southern America (a com-
plete list is available from the authors upon request),
including the Northern Apennine record. The exclu-
sion of the Southern American record is due to its
association with hypoxic or anoxic settings, favouring
preservation, but never encountered in other areas and
not evenly distributed in the Neogene. We matched in-
dividual specimens with sedimentary facies and time
interval, and listed the number of bones per skeleton,
ranging from the Aquitanian to the Calabrian (102
Northern Apennine specimens; 255 total specimens
from North America, Europe and Japan, with chro-
nostratigraphic data; 117 total specimens with data on
sedimentary facies). Given the extension of the stu-
dy to little known stratigraphic settings, stratigraphic
resolution was that of the stage for the Pliocene, or
Bullet-pointS aBStract
• The majority of cetacean fossils are in Zanclean and Piacenzian deposits.
• Cetacean fossils are preferentially found in offshore paleosettings.
• Pleistocene findings drop to a minimum, notwithstanding offshore strata are
well represented in the record.
• A taphonomic imprinting on the cetacean fossil record is hypothesised, con-
nected with a radiation of whale-bone consumers of modern type.
KeywordS:
Neogene;
Pliocene;
Cetaceans;
Taphonomy.
Corresponding author email: freschiales@gmail.com
How to cite: Dominici et al. (2018). Stratigraphic paleobiology of an evolutionary radiation: taphonomy and facies
distribution of cetaceans in the last 23 million years. Fossilia, Volume 2018: 15-17. https://doi.org/10.32774/FosRep-
Pal.20.1810.051517
Fossilia - Reports in Palaeontology
Dominici et al.
16 16
coarser for the Miocene and the Pleistocene.
reSultS
Our analysis highlights a strongly skewed distribution
of findings, both in space (seven facies bins) and time
(six time bins). The vast majority of fossils are found
in offshore mudstones (44%) and offshore sandstones
(28%), with condensed sections being particularly im-
portant in the Miocene. The lowest abundances are
recorded in shoreface sandstones, whereas complete-
ness, measured as the average number of bones per
skeleton, is higher in offshore mudstone and delta
sandstone facies, meaning that the two facies bear the
most complete skeletons. When plotting the number
of findings (N = 255) per time interval, standardizing
for the duration of each time bin (N/Ma), a slight in-
crease is recorded during the Miocene, a stepwise in-
crease at the passage upper Miocene-Zanclean, follow
by a Piacenzian peak and a dramatic drop at the Plei-
stocene, where the global number of skeletons falls to
a minimum (Fig. 1A). To understand the origin of this
bias, we analyzed (N= 131) the distribution of facies
in time and found a reasonably even distribution of
facies per time interval (Fig.1B). This pattern suggests
that the Piacenzian peak in biodiversity is in part an
artifact of the record.
diScuSSionS
In the Pliocene of Tuscany articulated specimens and
Fig. 1. A, Distribution (%) of the findings (N = 255) per time interval, standardized for the duration of each time interval (N/
Ma); B, Distribution of fossils (N = 131) and facies for time bin.
rather complete skeletons are associated with offshore
mudstones deposited at an estimated depth of 30–300
m, suggesting that very shallow and very deep (ba-
thyal) depths are generally unfavourable to the preser-
vation of bones (Dominici et al., 2018). An analogous
environmental control on the degree of articulation
and completeness of fossil cetacean skeletons is also
observed in the Pliocene of Emilia and Piedmont, po-
sitively correlated with offshore mudstones and bio-
calcarenite transgressive shell-beds, whereas no ske-
letons are associated with shoreface sandstones, and
rare occurrences in epibathyal mudstones (Freschi &
Cau, 2016).
Our study suggests that going from shallowest to de-
epest, the delta front processes allow skeletons to be
well-preserved due to high sedimentation rates that
quickly cover the carcasses that sequesters them before
they can be destroyed. In shoreface depths, carcasses
refloat after sinking, then become dismembered and
very frequently dispersed. On the contrary offsho-
re depths, carcasses sink and remain on the seafloor
because of a higher pressure that arrests abdominal
gas production, minimizes decomposition, and in the
absence of a specialized fauna, are not disassembled
before burial. Carcasses remain on the seafloor also
when they sink below the shelf break. However, here
they become the food for a complex and specialized
whale-fall fauna, characterized by bone-eating worms
and other taxa that finally destroy all bone material
(Smith et al., 2015). The dominance of offshore se-
diments in the Pliocene record explored here sugge-
17
taphonomy and facies distribution of cetaceans in the last 23 Ma
17
sts that the peak of Neogene biodiversity trend is in
part an artefact of a taphonomic influence. Finally, we
suggest that the sudden drop in the number of Plei-
stocene findings, notwithstanding the proper facies is
widespread in Italy, is also strongly affected by tapho-
nomic factors. The fact that this passage coincides
with the onset of the evolutionary radiation of very
large baleen whales and with their almost doubling in
maximum size (Marx et al., 2016) — which should
instead contribute a far better record — must have also
triggered the radiation of bone eaters, with the counte-
rintuitive effect of making extremely unlikely for ceta-
cean bones to become preserved. Modern worldwide
data confirm that even the largest whale skeletons are
rapidly destroyed at deep-water settings (see references
in Dominici et al., 2018).
concluSionS
In this work a large dataset of stratigraphic and tapho-
nomic global literature data of Neogene cetacean is
analysed (Southern America excluded). Abundance
and diversity are low during the Miocene, they peak
during the Pliocene, mainly in association with offsho-
re sediments, and drasticly drops in the Pleistocene,
irrespective of facies and notwithstanding the large
size attained during this epoch by cetaceans should be
correlated with a better record. The worsening of the
record during the Pleistocene may be related to a ra-
diation of bone-eaters, completely destroying carcas-
ses in a matter of years in modern bottoms. In gene-
ral, our study suggests a taphonomic imprinting in the
distribution of abundance and diversity of Neogene
cetaceans.
acKnowledgementS
We thank Raymond L. Bernor and an anonymous reviewer
for their useful suggestions.
referenceS
Berta A., Sumich J. & Kovacs K. (2015). Marine Mammals.
3rd Edition, Elsevier, 738 pp.
Dominici S., Danise S. & Benvenuti M. (2018). Pliocene
stratigraphic paleobiology in Tuscany and the fossil re-
cord of marine megafauna. Earth-Science Reviews, 176:
277–310.
Freschi A. & Cau S. (2016). Distribuzione geografica dei ce-
tacei fossili del Bacino Plio-pleistocenico di Castell’Ar-
quato (Piacenza). Parva Naturalia, 11: 47–59.
Freschi A., Cau S., Monegatti P. & Roveri M. (in press).
Chronostratigraphic distribution of cetaceans in the
Pliocene of Northern Apennines (Italy): palaeoecolo-
gical implications.
Marx G. M., Lambert O. & Uhen M. D. (2016). Cetacean Pa-
laeobiology. Topics in Paleobiology, Wiley-Blackwell,
336 pp.
Pimiento C., Griffin J. N., Clements C. F., Silvestro D., Vare-
la S., Uhen M. D. & Jaramillo C. (2017). The Pliocene
marine megafauna extinction and its impact on functio-
nal diversity. Nature ecology & evolution, 1 (8): 1100.
Smith C. R., Glover A. G., Treude T., Higgs N. D., Amon
D. J. (2015). Whale-fall ecosystems: recent insights into
ecology, paleoecology, and evolution. Annual Review of
Marine Science, 7: 571-596.
Steeman M. E., Hebsgaad M. B., Fordyce R. E., Ho S. Y.
W., Rabosky D. L., Nielsen R., Rahbek C., Glenner H.,
Sorensen M. V., & Willerslev E. (2009). Radiation of
extant cetaceans driven by restructuring of the oceans.
Systematic Biology, 58: 573-585.
Manuscript received 27 July 2018
Received after revision 29 September 2018
Accepted 1 October 2018
Fossilia, Volume 2018: 19-22
Early Miocene
Megacricetodon
and
Democricetodon
(Cricetidae, Rodentia) from the
Vallès-Penedès Basin (Catalonia)
Sílvia Jovells-Vaqué & Isaac Casanovas-Vilar
Institut Català de Paleontologia Miquel Crusafont, Universitat Autònoma de Barcelona, Cerdanyola del Vallès, Spain. silvia.jovells@icp.
cat; isaac.casanovas@icp.cat
introduction
The cricetodontine genera Democricetodon Fahlbusch,
1964 and Megacricetodon Fahlbusch, 1964 are the first
modern cricetids to disperse into Europe. Democriceto-
don is first recorded during the late MN3, whereas Me-
gacricetodon does not appear until MN4 (Agustí et al.,
2001; Hilgen et al., 2012). These cricetids dispersed
after the extinction of the archaic cricetids Pseudocri-
cetodon (Thaler, 1969) and Eucricetodon (Thaler, 1966),
which had characterized the Oligocene, and quickly
became dominant in the rodent faunas. Megacricetodon
and Democricetodon have been widely used in biostrati-
graphical and biochronological scales for the early and
middle Miocene all along Europe, the occurrence of
their different species being the basis for the definition
of high-resolution local biozones (e.g. Van der Meulen
et al., 2012; Kälin & Kempft, 2009; Prieto & Rummel,
2016). In this work, we review the early Miocene re-
cord of Megacricetodon and Democricetodon in the Val-
lès-Penedès Basin (Catalonia, NE Iberian Peninsula).
The early Miocene part of the record of this basin has
been little studied in comparison with middle to late
Miocene one.
materialS and methodS
Between 2011 and 2017 the early Miocene outcrops
were systematically surveyed resulting in the discovery
of several new localities (Casanovas-Vilar et al., 2016).
In addition, some of the classical sites were sampled
using modern methods, including screen-washing,
thus allowing for the recovery of remarkably rich mi-
crovertebrate samples (Jovells-Vaqué et al., 2018). The
described material is housed at the Institut Català de
Paleontologia Miquel Crusafont in Sabadell (Barcelo-
na, Spain).
SyStematic palaeontology
Order Rodentia Bowdich, 1821
Family Cricetidae Fischer, 1817
Genus Democricetodon Fahlbusch, 1964
Democricetodon hispanicus Freudenthal, 1967
Figs. 1A-D
The m1 present a characteristic bean-shaped antero-
conid. The anterior valleys of the lower teeth are clo-
sed and reduced. The mesolophid varies from long to
short in m1 and m2 (Figs. 1B-C) but is completely ab-
sent in the m3. This ridge is more frequently short to
medium length, being generally longer in the m1 than
m2.The sinusid is wide and slightly proverse in the m1
and m2, while in m3 is moderately retroverse. The M1
show a simple round anterocone (Fig.1A). The meso-
loph varies from short to medium length in M1 and
M2. The anterosinus is reduced in M2 and M3 due to
the reduction of the anteroloph.
Democricetodon cf. decipiens Freudenthal & Daams,
1988
In Vilobí del Penedès and les Escletxes del Papiol sites
besides D. hispanicus a second, larger species of Demo-
cricetodon is represented by a few specimens. These fit
within the size range of Democricetodon decipiens. In ad-
dition they show some of the diagnostic traits of this
species such as the presence of shorter mesolophs/ids
than D.hispanicus.
Democricetodon cf. moralesi Van der Meulen et al., 2003
Fig. 1E
Bullet-pointS aBStract
• Democricetodon and Megacricetodon species are present during the early Miocene
in the Vallès-Penedès Basin.
• The first occurrence of both genera Democricetodon and Megacricetodon in the
Basin mark the beginning of the MN4.
• These species are key for stablish a high-resolution local biozonation and corre-
lated to other European Basins.
KeywordS:
early Miocene;
Democricetodon;
Megacricetodon;
Biostratigraphy;
Iberian Peninsula.
Corresponding author email: silvia.jovells@icp.cat
How to cite: Jovells-Vaqué & Casanova-Villar (2018). Early Miocene Megacricetodon and Democricetodon (Cricetidae,
Rodentia) from the Vallès-Penedès Basin (Catalonia). Fossilia, Volume 2018: 19-22. https://doi.org/10.32774/FosRep-
Pal.20.1810.061922
Fossilia - Reports in Palaeontology
20
An even larger-sized Democricetodon species is present
at els Casots and la Riera del Morral sites. At els Ca-
sots it cooccurs with D. hispanicus, whereas at Riera
del Morral it represents the only recovered species.
This species is larger than both D. hispanicus and D. de-
cipiens, the few recovered specimens fitting within the
range size of D. moralesi. The recovered M2 share with
D. moralesi the presence of a double protolophule in
the M2 as well as a short to medium-sized mesolophs/
ids.
Democricetodon cf. gracilis Fahlbusch, 1964
Finally, yet another species is present at some sites (els
Casots, Vilobí del Penedès, Sant Mamet), although it
is represented by very few specimens. These are distin-
guished because of their small size, below the size ran-
ge of D. hispanicus.
Genus Megacricetodon Fahlbusch, 1964
Megacricetodon primitivus (Freudenthal, 1963)
Figs. 1F-J
Small-sized species of Megacricetodon. The m1 present
a simple rounded anteroconid. The lingual antero-
lophid is reduced, thus resulting in a narrow antero-
sinusid. The mesolophid ranges from long to short,
but is more frequently short to medium lenght. The
M1 (Fig. 1F) shows a deeply split anterocone with a
well-defined platform-like anterior cingulum on its
base. The protolophule is simple in the M1 and M2.
The mesoloph ranges from long to medium length.
diScuSSionS and concluSion
The genera Democricetodon and Megacricetodon first ap-
pear simultaneously in MN4 sites of the Vallès-Pened-
ès Basin. In other European basins, such as the Cala-
tayud-Montalbán Basin or the Swiss Molasse Basin,
Democricetodon is recorded earlier, during the late MN3
(Kälin & Kempf, 2009; Van der Meulen et al., 2012).
The situation seen in the Vallès-Penedès might be at-
tributable to the lack of late MN3 sites (correlated to
zone B of Calatayud-Montalbán Basin, the type area
of the Aragonian mammal age). The species identified
in the Vallès-Penedès sites are also recorded in the Ca-
latayud Montalbán Basin, although they are not coe-
taneous, D. hispanicus being restricted to local zone B
and M. primitivus first occurring in local subzone Ca
(Van der Meulen et al., 2012). Local subzone Ca is
Fig. 1. Scanning electron microscope (SEM) micrographs of the Cricetidae from els Casots site. Modified from Jovells-Vaqué et
al. 2017. Democricetodon hispanicus Freudenthal, 1967 – A, IPS 45008, right M1-M3 (reversed); B, IPS 45049, right m1 (reversed);
C, IPS 45059, right m2 (reversed); D, IPS 19473, left m3. Democricetodon cf. moralesi Van der Meulen et al., 2003– E, IPS 45052,
left m1. Megacricetodon primitivus (Freudenthal, 1963) – F, IPS 44939 right M1 (reversed); G, IPS 44961, left M2; H, IPS 19479
left M3; I, IPS 44950, right m1-m2 (reversed); J, IPS:44992 right m3 (reserved).
Jovells-Vaqué & Casanovas-Vilar
21
also characterized by the coexistence of the eomyids
Ligerimys florancei (Stehlin & Schaub, 1951) and Lige-
rimys ellipticus (Daams, 1976), which also co-occur in
all the Vallès-Penedès sites except els Casots, Vilobí
del Penedès and Sant Mamet. These could indicate
that the latter localities might be somewhat younger.
The fact that they have delivered additional, larger-si-
zed Democricetodon species, Democricetodon cf. decipiens
and Democricetodon cf. moralesi, supports this interpre-
tation. D. decipiens characterizes subzone Ca in the
Aragonian type area, whereas D. moralesi is diagnostic
of zone Cb. In the Vallès-Penedès Basin the cricetid
succession is not the same, D. hispanicus persists for a
longer time (well into zone C) and a clear distinction
between subzones Ca and Cb on the basis of the cri-
cetid species is not possible. On the other hand, the
small-sized Democricetodon cf. gracilis is not present
in the Calatayud-Montalbán Basin, being characteri-
stic of Central Europe (Swiss and Bavarian Molasse
basins; see Abdul Aziz et al., 2008; Kälin & Kempf,
2009). Therefore, the Vallès-Penedès cricetid record
includes taxa mostly showing affinities with other Ibe-
rian basins as well as a minor proportion of forms that
evidence connections with Central Europe.
acKnowledgementS
This research has been funded by the Agencia Estatal de In-
vestigación from the Spanish Ministerio de Economía, In-
dústria y Competitividad and the Agencia Estatal de Investi-
gación (AEI) from Spain/European Regional Development
Fund of the European Union (project CG2017-82654-P,
CG2016-76431-P, RYC-2013-12470 research contract to
ICV), by the Agència de Gestió d’Ajuts Universitaris i de
Reserca of the Generalitat de Catalunya (2018 FI B1 00201
pre-doctoral grant to SJV), and by the CERCA programme
of the Generalitat de Catalunya (project 2014/100584) and
the National Geographic Society (grant number 9645-15).
This work has been performed by researchers from the con-
solidated research group “Neogene and Quaternary Verte-
brate Paleobiodiversity (NQVP)” (2017 SGR 86 GRC) of
the Generalitat de Catalunya.
referenceS
Abdul Aziz H., Böhme M., Rocholl A., Zwing A., Prieto J.,
Wijbrans J. R., Heissig K. & Bactadse V. (2008). Inte-
grated stratigraphy and 40Ar/39Ar chronology of Early
to Middle Miocene Upper Freshwater Molasse in ea-
stern Bavaria (Germany). International Journal of Earth
Sciences, 97:115–134.
Agustí J., Cabrera L., Garcés M., Krijgsman W., Oms O.
& Parés J. M. (2001). A calibrated mammal scale for
the Neogene of Western Europe. State of the art. Earth
Science Reviews, 52: 247–260.
Bowdich T. E. (1821). An Analysis of the Natural Classifica-
tions of Mammalia for the Use of Students and Travel-
lers. Ed: J. Smith. Paris, 115 pp.
Casanovas-Vilar I., Madern A., Alba D. M., Cabrera L.,
García-Paredes I., Van den Hoek Ostende LW., DeMi-
guel D., Robles JM., Furió M., van Dam JA., Garcés
M., Angelone C. & Moyà-Solà S. (2016). The Miocene
mammal record of the Vallès-Penedès Basin (Catalo-
nia). Comptes Rendus Palevol, 15:791–812.
Daams R. (1976). Miocene rodents (Mammalia) from Ce-
tina de Aragón (prov. Zaragoza) and Buñol (prov. Va-
lencia), Spain. Proceedings of the Koninklijke Akademie van
Wetenschappen B, 79: 152–182.
Fahlbusch V. (1964). Die Cricetiden (Mamm.) Der Oberen
Sübwasser-Molasse Bayerns. Bayerische Akademie der
Wissenschaften Mathematisch-Natur Wissenschaftliche Klas-
se Abhandlungen, 118: 4–136.
Fischer W. (1817). Adversaria zoologica. Mémories de la So-
ciété Impériale des Naturalistes de Moscou, 5: 357-471
Freudenthal M. (1963). Entwicklungsstufen der miozänen
Cricetodontinae (Mammlia, Rodentia) Mittlespaniens
und ihre stratigraphische Bedeutung [Succession of
Miocene Cridetodontinae (Mammalia, Rodentia) in
central Spain and their stratigraphic significance]. Be-
aufortia, 10:51–157.
Freudenthal M. (1967). On the Mammalian Fauna of the
Hipparion-Beds in the Calatayud-Teruel Basin (Prov.
Zaragoza, Spain). Part III. Democricetodon and Ro-
tundomys (Rodentia). Proceedings of the Koninklijke Aka-
demie van Wetenschappen, 70 (3): 298–315.
Freudenthal M. & Daams R. (1988). Cricetidae (Rodentia)
from the Type-Aragonian; the Genera Democriceto-
don, Fahlbuschia, Pseudofahlbuschia Nov. Gen., and
Renzimys. In: Biostratigraphy and Paleoecology of
the Neogene Micromammalian Faunas from the Ca-
latayud-Teruel Basin (Spain). Freudenthal M. (Eds).
Scripta Geologica, Special Issue 1: 133–252.
Hilgen F. J., Lourens L. J. & Van Dam J. A. (2012). The
Neogene period. In Gradstein FM, Ogg JG, Schmitz M
& Ogg G. (eds), The geologic time scale. Elsevier, Am-
sterdam: 923–978.
Jovells-Vaqué S., Ginestí M. & Casanovas-Vilar I. (2017).
Cricetidae (Rodentia, Mammalia) from the early Mio-
cene site of els Casots (Vallès-Penedès Basin, Catalo-
nia). Fossil Imprint, 73: 141-154.
Jovells-Vaqué S., García-Paredes I., Furió M., Angelone C.,
Van den Hoek Ostende LW., Berrocal Barberà M., De-
Miguel D., Madurell-Malapeira J. & Casanovas-Vilar I.
(2018). Les Cases de la Valenciana, a new early Mio-
cene small-mammal locality from the Vallès-Penedès
Basin (Catalonia, Spain). Historical Biology, 30: 404-421.
Kälin D. & Kempf O. (2009). High-resolution stratigraphy
from the continental record of the Middle Miocene
Northern Alpine Foreland Basin of Switzerland. Neu-
es Jahrbuch für Geologie und Paläontologie - Abhandlungen,
254: 177–235.
Prieto J. & Rummel M. (2016). Some considerations about
the small-mammal evolution in South Germany, with
emphasis on late Burdigalian-earliest Tortonian (Mio-
cene) cricetid rodents. Comptes Rendus Palevol, 15 : 837-
854.
Stehin H. G. & Schaub S. (1951). Die Trigonodontie de
simplicidentaten Nager. Schweizerische Palaontologische
Abhandlung, 67: 1-385.
Thaler L. (1966). Les rongeurs fossiles du Bas-Languedoc
dans leurs rapports avec I’histoire des faunes et la stra-
tigraphie du Tertiaire d’Europe. Memoires du Museum
National d’Histoire Naturelle, 17: 295 pp.
Thaler L. (1969). Rongeurs nouveaux de l’Oligocène moyen
d’Espagne. Paleovertebrata, 2: 191-207
Van der Meulen A. J., Pélaez-Campomanes P. & Daams R.
(2003). Revision of Medium-Sized Cricetidae from the
Miocene of the Daroca-Villafeliche Area in The Cala-
tayud-Teruel Basin (Zaragoza, Spain). Coloquios de Pale-
ontologia, 1: 385–441.
Megacricetodon
and
Democricetodon
from the Vallès-Penedès Basin
22
Manuscript received 10 July 2018
Received after revision 2 October 2018
Accepted 3 October 2018
Van der Meulen A. J., García-Paredes I., Álvarez-Sierra M.
A., Van den Hoek Ostende L. W., Hordijk K., Oliver
A. & Peláez-Campomanes P. (2012). Updated Arago-
nian biostratigraphy: small mammal distribution and
its implications for the Miocene European chronology.
Geologica Acta, 10: 159–179.
Jovells-Vaqué & Casanovas-Vilar
Fossilia, Volume 2018: 23-25
Histological study of sauropod dinosaur bones
from the historic Upper Jurassic Howe Quarry
(Wyoming, USA): determination of an age range for
every specimen
Jacopo Moretti1, Emanuel Tschopp2, Daniel Barta2, Katja Waskow3 & Mark Norell2
1 Dipartimento di Scienze Chimiche e Geologiche, Università di Modena e Reggio Emilia, Via Giuseppe Campi 103, 41125 Modena, Italy;
jachi.moretti@hotmail.it.
2American Museum of Natural History, Central Park West & 79° street, 10024 New York, NY, USA.
3Steinmann Institute for Geology, Mineralogy, and Paleontology, University of Bonn, Nussallee 8, 53113 Bonn, Germany.
introduction
The Howe Quarry, in the vicinity of Shell (Wyoming,
USA), is one of the richest occurrences of sauropod
bones in the Upper Jurassic Morrison Formation (Fo-
ster, 2007). Excavations in 1934 by the American Mu-
seum of Natural History (AMNH, New York, USA)
(Brown, 1935; Bird, 1985) and in 1990 and 1991 by
the Sauriermuseum Aathal (Switzerland) (Ayer, 2000)
produced approximately 3000-4000 bones. The majo-
rity of the recovered specimens are sauropod dinosau-
rs, most of them belonging to the clade Diplodocidae.
The aim of this study was to analyze sauropod bones
collected from the Howe Quarry, both morphological-
ly and histologically.
materialS and methodS
It was decided to focus on long bones and ribs, as these
are the most useful bones to identify histological featu-
res. Long bones, developing an isometric growth pat-
tern, can be analyzed to determine the size and mass
of the specimens, whereas ribs, which do not have an
isometric growth pattern, make it easier to identify
histological features (Tschopp & Mateus, 2013). The
fossils were firstly measured and scanned to obtain a
digital database. After that, they were sectioned and
then studied under the optical microscope to identify
histological characteristics such as LAGs (Lines of Ar-
rested Growth), EFS (External Fundamental System)
and osteons. LAGs are growth lines representing the
former surface of the bone. The EFS is a structure visi-
ble under the surface of the bone, made of very closely
spaced LAGs, which indicates the skeletal maturity of
the specimen (Fig. 1).
The eight specimens analyzed by this study corre-
spond to seven ribs (AMNH FARB 30786, AMNH
FARB 30791, AMNH FARB 30901, AMNH FARB
33141, AMNH FARB 33144, AMNH FARB 33145,
AMNH FARB 33147) and one fibula (AMNH FARB
30785). Following two previous studies on the same
subject, histological data were gathered from the ribs
counting LAGs, generations of secondary osteons and
searching for the presence of an EFS.
reSultS and diScuSSionS
At least three different ontogenetic stages could be
identified among the sampled specimens: 1) early juve-
niles; 2) sexually but not skeletally mature adults; and
3) skeletally mature adults. Curves showing the deve-
lopment of LAGs, depending on their distance from
each other and their distance from the center of the
bone, were produced and compared showing, in some
cases, a perfect overlap of the data, resulting in curves
from different specimens following an identical trend.
The results witness a diversified fossil assemblage cha-
racterized by the presence of individuals with different
ages and ontogenetic development (Figs. 2-3).
Resulting histological data were then compared with
Waskow & Sander (2014) to obtain more information
Bullet-pointS aBStract
• Morphological and Histological study on sauropod bones from the Howe
Quarry with the determination of an age range and ontogenetic stage for
every specimen.
• Eight specimens studied, one fibula and seven ribs: three skeletally mature,
three sexually mature, one juvenile and one undetermined.
• Useful and generally applicable method without a taxonomical assignment.
KeywordS:
Howe Quarry;
sauropods;
bone histology;
age range;
ontogenetic stage.
Corresponding author email: jachi.moretti@hotmail.it
How to cite: Moretti et al. (2018). Histological study of sauropod dinosaur bones from the historic Upper Jurassic Howe
Quarry (Wyoming, USA): determination of an age range for every specimen. Fossilia, Volume 2018: 23-25. https://doi.
org/10.32774/FosRepPal.20.1810.092325
Fossilia - Reports in Palaeontology
Moretti et al.
24
on the degree of skeletal maturity of every specimen.
The only specimen which could be associated with a
very juvenile stage is AMNH FARB 33144. It does not
present any LAG or secondary osteons. These histolo-
gical characteristics are in accordance with the maxi-
mum diameter of the rib, which is the smallest among
the studied specimens.
Three specimens, AMNH FARB 30786, AMNH
FARB 30791 and AMNH FARB 33145, were in-
terpreted as sexually mature, an intermediate stage
between juveniles and skeletally mature individuals.
They all have LAGs and osteons but do not show any
sign of an EFS.
The last three specimens, AMNH FARB 30901,
AMNH FARB 33141 and AMNH FARB 33147, were
identified as skeletally mature for the presence of an
EFS.
The fibula AMNH FARB 30785 did not give any hi-
Fig. 1. Resume of images and data of AMNH FARB 30901. A, 3D scan of the dorsal rib (the red line indicates sectioning of
following frames). B, thin section (and detail on the right) illustrating LAGs (in red), the EFS is only visible under optical micro-
scope, white holes represent osteons. C, data gathered from this specimen.
Fig. 2. General graphic representing the distances of LAGs
from the center of the rib in all the studied specimens.
Fig. 3. General graphic representing the distances between
LAGs in all the studied specimens.
25
stological information, therefore it was not possible to
classify it in terms of ontogenetic stage.
concluSionS
In the end, it was possible to divide all the studied spe-
cimens in three age ranges and determine the onto-
genetic stage for each one of them. It is also possible
to say that there are no evidences of dwarf specimens
in this association, because the smallest specimen is
also the most juvenile. One of the most interesting
characteristics of this method is that it was applied
without considering the taxonomy of the animal, whi-
ch is usually difficult to determine from a single bone
specimen.
acKnowledgementS
Amy Davidson, Verne Lee, Mick Ellison, Carl Mehling,
Judy Galkin (all of the American Museum of Natural Hi-
story) and Cary Woodruff (University of Toronto).Daniel
Barta is supported by a Richard Gilder Graduate School Fel-
lowship. E. Tschopp is supported by a Theodore Roosevelt
Memorial Fund and Division of Paleontology Postdoctoral
Fellowship. Massimo Delfino (University of Turin) and An-
nalisa Ferretti (University of Modena and Reggio Emilia).
referenceS
Ayer J. (2000). The Howe Ranch Dinosaurs. Sauriermu-
seum Aathal, Aathal, Switzerland.
Bird R. T. (1985). Bones for Barnum Brown, adventures of a
dinosaur hunter. Texan Christian University Press, Fort
Worth, U.S.A.
Brown B. (1935). Sinclair Dinosaur Expedition, 1934. Eds.
Natural History, 1934, New York City (NY), 3-16.
Foster J. (2007). Jurassic West: The Dinosaurs of the Mor-
rison Formation and Their World, Indiana University
Press, Bloomington, U.S.A.
Tschopp E. & Mateus O. (2013). The skull and neck of a
new flagellicaudatan sauropod from the Morrison For-
mation and its implication for the evolution and onto-
geny of diplodocid dinosaurs. Journal of Systematic Pala-
eontology, 11: 853–888.
Waskow K. & Sander P.M. (2014). Growth record and hi-
stological variation in the dorsal ribs of Camarasaurus
sp. (Sauropoda). Journal of Vertebrate Paleontology, 34:
852-869.
Manuscript received 14 July 2018
Received after revision 21 September 2018
Accepted 2 October 2018
Histology of sauropod bones: age range determination
Fossilia, Volume 2018: 27-32
Evolutionary history of Rhinocerotina
(Mammalia, Perissodactyla)
Luca Pandolfi
Dipartimento di Scienze della Terra, Università degli Studi di Firenze 50121, Italy; luca.pandolfi@unifi.it
introduction
The tribe Rhinocerotina Gray, 1821 (sensu Antoine,
2002) includes the extant rhinoceroses that are cur-
rently distributed in limited areas of SouthAsia and
Africa.
The tribe probably originates from Late Oligocene re-
presentatives of Teleoceratini, but this hypothesis ne-
eds to be better supported by a comprehensive revision
of the latter taxon. Rhinocerotina is considered a mo-
nophyletic group (Antoine, 2002) supported by seve-
ral synapomorphies such as the presence of a median
frontal horn, enlarged rostral end of the nasals, trend
towards the ossification of the nasal septum (Antoine,
2002; Antoine et al., 2003).
materialS and methodS
This note is based on specimens, from both original
observations and published material, referred to the 5
extant rhinocerotine species and several extinct rhino-
cerotine taxa (Fig. 1; Tab. 1), that occur in the Neoge-
ne and Quaternary fossil record of Eurasia and Africa
(Fig. 1). The species list and the list of institutions,
where the observed specimens are preserved, are re-
ported in Tab. 1.
Institutional abbreviations
BSPG, Bayerische Staatssammlung für Paläontologie
und Geologie, Munich, Germany; HNHM, Hunga-
rian Natural History Museum, Budapest, Hungary;
IGF, Istituto Geo-Paleontologico di Firenze, Floren-
ce, Italy; IRSNB, Institut royal des Sciences naturel-
les de Belgique, Bruxelles, Belgium; IVPP, Institute
of Vertebrate Paleontology and Paleoanthropology,
Beijing, China; MAFI, Magyar Földtani és Geofizikai
Intézet (Geological and Geophysical Institute of Hun-
gary), Budapest, Hungary; MfN, Museum für Na-
turkunde, Berlin, Germany; MGGC, Museo di Ge-
ologia Giovanni Capellini, Bologna, Italy; MPGPD,
Museo di Geologia e Paleontologia, Padua, Italy;
MNCN, Museo Nacional de Ciencias Naturales, Ma-
drid, Spain; MNHN, Musèum National d’Histoire
naturelle, Paris, France; MPP, Museo Paleontologico
Parmense, Parma, Italy; MPUR, Museo Paleontolo-
gico dell’Università di Roma, Rome, Italy; MSNAF,
Museo di Storia Naturale, Accademia dei Fisiocritici,
Siena, Italy; MZF, Museo di Storia Naturale, sezione
di Zoologia, Florence, Italy; NHMUK, Natural Hi-
story Museum, London, United Kingdom; NHMW,
Naturhistorisches Museum, Wien, Austria; NMB,
Naturhistorisches Museum, Basel, Switzerland; SMF,
Senckenberg Naturmuseum, Frankfurt, Germany;
ZSM, Zoologische Staatssammlung, Munich, Ger-
many.
Genus abbreviations
C. = Ceratotherium; Co. = Coelodonta; D. = Dicerorhinus;
Di. = Diceros; Dh. = Dihoplus; G. = Gaindatherium; L. =
Lartetotherium; P. = Paradiceros; R. = Rhinoceros; Ru. =
Rusingaceros; S. = Stephanorhinus.
diScuSSionS and concluSionS
The earliest European representative of Rhinocero-
tina is Lartetotherium sansaniense (Lartet in Laurillard,
1848), which occurred during the MN4 (early Mioce-
ne). Rusingaceros leakeyi (Hooijer, 1966) and “Diceros”
australis Guérin, 2000 (both early Miocene in age) are
documented in African Continent, whereas the tribe
occurred in South Asia as soon as the early Mioce-
ne, with Gaindatherium cf. browni and cf. Rhinoceros sp.
(Bugti Hills, Pakistan; Antoine et al., 2013) (Fig. 1).
The maximum geographic distribution of the tribe oc-
curred in the latest Miocene and the Plio-Pleistocene,
when all aceratheriines and teleoceratines went ex-
tinct and elasmothere’s diversity drastically reduced.
Four latest Miocene Rhinocerotina were certainly
Bullet-pointS aBStract
• The earliest representatives of the group occurred during the Early Miocene.
• The group is well documented during the Late Miocene in Western Eurasia
and Africa.
• High species diversity occurred in Eurasia during the Pleistocene.
• The fossil records of Rhinoceros and Dicerorhinus seems to be confined in the
Indian subcontinent and East and South-East Asia.
KeywordS:
Rhinocerotina;
evolution;
paleobiogeography;
Neogene;
Quaternary.
How to cite: Pandolfi (2018). Evolutionary history of Rhinocerotina (Mammalia, Perissodactyla). Fossilia, Volume 2018:
27-32. https://doi.org/10.32774/FosRepPal.20.1810.102732
Fossilia - Reports in Palaeontology
Pandolfi
28
Fig. 1. Distribution of extant and fossil species of subtribe Rhinocerotina. The species are rougly listed according to their occurrences for each epoch and for each considered area.
Evolutionary history of Rhinocerotina 29
Species Direct Observation References
"D.” steinheimensis NHMW Guérin, 1980; Heissig, 1999
“Di.” australis -Guérin, 2000; Geraads, 2010
“S.” miguelcrusafonti MNCN Guérin, 1980; Cerdeño, 1992
C. douariense MfN, NHMUK Guérin, 1966; Giaourtsakis et al., 2009; Geraads, 2010;
C. mauritanicum -Geraads, 2005; 2010
C. neumayri NHMUK, NHMW, NMB Guérin, 1980; Geraads, 1988; 2005; Giaourtsakis,
2009; Antoine & Saraç, 2005; Giaourtsakis et al., 2006
C. simum ssp. IRSNB, MfN, NHMUK, NMB Guérin, 1980
C.? primaevum -Arambourg, 1959; Geraads, 2010
Co. antiquitatis HNHM, IGF, MAFI, MfN, MGGC, MPGPD,
MPUR, NHMUK, NHMW, NMB
Guérin, 1980
Co. nihowanensis -Deng, 2002
Co. thibetana -Deng et al., 2011
Co. tologojiensis -Belyaeva, 1966; Kahlke & Lacombat, 2008
Co. antiquitatis -Guérin, 1980
D. cixianensis -Chen & Wu, 1976
D. gwebinensis -Zin-Maung-Maung-Thein et al., 2008
D. sumatrensis MNHN, MZF, NHMUK, NMB Guérin, 1980
Di. bicornis ssp. MfN, MNHN, NHMUK, NMB, SMF, ZSM Guérin, 1980
Di. gansuensis -Deng & Qiu, 2007
Di. praecox MfN, NHMUK Hooijer & Patterson, 1972; Guérin, 1987; Geraads,
2010
Dh. megarhinus BSPG, IGF, MGGC, MPP, MSNAF, NHMUK, NMB Guérin, 1980; Pandolfi et al., 2015 and references
therein
Dh. pikermiensis NHMUK, NHMW Geraads, 1988; Giaourtsakis, 2009; Giaourtsakis et al.,
2006
Dh. schleiermacheri HNHM, MAFI, MfN, MNCN, NHMUK, NHMW,
NMB
Guérin, 1980; Cerdeño, 1992
G. browni -Colbert, 1934; Khan et al., 2014
G. vidali -Heissig, 1972; Khan et al., 2014
L. sansaniense MfN, MNCN, NMB Guérin, 1980; Cerdeño, 1996; Heissig, 2012
P. mukirii NHMUK Hooijer, 1968; Geraads, 2010; Guérin, 2011
R. platyrhinus NHMUK, Falconer & Cautley, 1846; Colbert, 1935; Khan, 1971;
Pandolfi & Maiorino, 2016
R. sinensis -Matthew & Granger, 1929; Colbert & Hoojier, 1953
R. sivalensis NHMUK Falconer & Cautley, 1846; Colbert, 1935; Pandolfi &
Maiorino, 2016
R. sondaicus NHMUK, NMB Guérin, 1980
R. unicornis NHMUK, NMB Guérin, 1980
Ru. leakeyi NHMUK Hooijer, 1966; Geraads, 2010
S. etruscus IGF, MGGC, MNCN, MNHN, MPGPD, MPUR,
MSNAF, NHMUK, NMB
Guérin, 1980
S. hemitoechus IGF, MNCN, MPUR, NHMUK Guérin, 1980
S. hundsheimensis MNHN, NHMW, NHMUK Guérin, 1980
S. jeanvireti IGF, HNHM, MGGC, NMB Guérin, 1972; 1980
S. kirchbergensis MfN, MNHN, MPUR, NHMUK, NMB Guérin, 1980; Handa & Pandolfi, 2016
S. lantianensis IVPP Tong, 2012
S. yunchuchenensis IVPP Chow, 1963; Tong, 2012
S.? africanus -Arambourg, 1970; Geraads, 2010
Tab. 1. List of the species included within the subtribe Rhinocerotina and sources for the considered cranial and dental mate-
rial. For the genus abbreviation, check the Materials and Methods section.
identified in Eurasia: Dihoplus schleiermacheri (Kaup,
1832) (which occurred from MN 9 to MN 12 at seve-
ral central and western European localities), Dihoplus
pikermiensis (Toula, 1906) (which occurred in the latest
Vallesian and Turolian deposits, MN 10-MN 13, of
the Balkan Peninsula and Turkey), Dihoplus megarhi-
nus (de Christol, 1832) (recently reported from several
latest Miocene faunas, MN 12-MN 13, of Hungary,
Italy, Ukraine, and China), and Ceratotherium neumayri
(Osborn, 1900) (reported from several fossiliferous
localities, MN 10-MN 13, of the Balkan Peninsula,
Caucasus, Anatolia and Iran). Late Miocene African
localities which yielded remains of rhinoceros are re-
latively scarce and often the rhinoceros material was
only identified at the generic level (Ceratotherium Gray,
1868 or Diceros Gray, 1821; Geraads, 2010) or as Para-
diceros sp., cf. Ceratotherium sp. or cf. Paradiceros mukirii
Hooijer, 1968. According to Guérin (2011) a taxon
closely related with the Balkano-Iranian species C.
neumayri (Diceros cf. pachygnatus in Guérin, 2011) was
present at Aragai and Ngetabkwony (Kenya, around 6
Ma and 5-4.5 Ma respectively). Ceratotherium douarien-
se (Guérin, 1966) (= Diceros douariense) was established
from a partial skull with associated mandible from
Douaria (Tunisia, around 7 Ma) but it is doubtfully di-
stinct from C. neumayri (Geraads, 2010). The record of
C. douariense from the Middle Awash (Ethiopia, latest
Miocene; Giaourtsakis et al., 2009) was reported as
Diceros? sp. by Geraads (2010: Tab. 34.I) but without
any detailed explanation. Ceratotherium? primaevum
(Arambourg, 1959) (= Dicerorhinus primaevus Aram-
bourg, 1959) was named on a partial juvenile skull and
associated remains collected at Bou Hanifia (Algeria,
around 10 Ma).
During Plio-Pleistocene, the genera Stephanorhinus
Kretzoi, 1942 and Coelodonta Bronn, 1831 were wide-
spread in Eurasia (Fig. 1), but they never occurred in
the Indian Subcontinent and in the South-East Asia
(e.g., Thailand, Malaysia, Indonesia). Coelodonta and
Stephanorhinus are generally considered sister groups,
but Heissig (1981) suggested that any similarities
between these two taxa could be related to convergen-
ce.
The earliest representatives of the genus Stephanorhi-
nus are from the Pliocene of Europe. At least 7 species
can be included within this genus: Stephanorhinus kirch-
bergensis (Jäger, 1839) (Early Pleistocene to latest Plei-
stocene; Eurasia), Stephanorhinus etruscus (Falconer,
1868) (late Pliocene to the early Middle Pleistocene;
Europe), Stephanorhinus hemitoechus (Falconer, 1859)
(Middle Pleistocene to the latest Pleistocene; Eura-
sia and North Africa), Stephanorhinus hundsheimensis
(Toula, 1902) (latest Early Pleistocene to the Middle
Pleistocene; Europe), Stephanorhinus yunchuchenen-
sis (Chow, 1963) (latest Early Pleistocene; Eastern
China), Stephanorhinus jeanvireti (Guérin, 1972) (Late
Pliocene to earliest Pleistocene; Europe), and Stepha-
norhinus lantianensis (Hu & Qi, 1978) (latest Early Plei-
stocene; Eastern China). “Stephanorhinus” miguelcrusa-
fonti (Guérin & Santafé-Llopis, 1978) (Early Pliocene;
Western Europe) is provisionally retained within this
genus, whereas Stephanorhinus? africanus (Arambourg,
1970) (early to middle Pliocene; Tunisia and Chad)
has an uncertain taxonomic position. Heissig (1999)
suggested a paraphyly of Stephanorhinus, considering
two evolutionary lineages, one from Dihoplus schleier-
macheri to S. jeanvireti through “D.” megarhinus, and the
other one from Dihoplus pikermiensis (= Stephanorhinus
pikermiensis in Heissig, 1999) to S. etruscus and other
species. However, the debate on the phylogenetic rela-
tionships within this group of rhinoceroses is far from
being settled by consensus.
Coelodonta evolved in Central Asia since the middle
Pliocene (with the species Coelodonta thibetana Deng et
al., 2011 from the Tibetan Plateau) and reached We-
stern Europe only during the Middle Pleistocene; wi-
thin this genus the major apomorphies were acquired
by Coelodonta antiquitatis (Blumenbach, 1799). During
the last glacial maximum, the woolly rhino was pre-
sent from Iberian Peninsula to Siberia and from Scot-
land to Greece.
The genera Ceratotherium [with C. mauritanicum (Po-
mel, 1888) and C. simum (Burchell, 1817)] and Dice-
ros (with D. praecox (Hooijer & Patterson, 1872) and
D. bicornis (Linneaus, 1758)) were widespread throu-
ghout Africa during the Plio-Pleistocene. C. mauri-
tanicum was present from the Late Pliocene to Late
Pleistocene, in particular in North Africa, and it was
probably the ancestor of C. simum (Geraads, 2010),
which occurred for the first time in East Africa during
the Early Pleistocene. Contrary to C. simum, D. bicornis
has never been recorded North of the Sahara desert.
The genus Rhinoceros Linneaus, 1758 was mainly
limited in the Indian Subcontinent and some areas of
Southeastern Asia during the Pleistocene and Holo-
cene (Fig. 1). Rhinoceros sivalensis Falconer & Cautley,
1846 and Rhinoceros platyrhinus Falconer & Cautley,
1846 co-occurred in Southern Himalayas (Upper
Siwaliks). Rhinoceros unicornis Linneaus, 1758, current-
ly is represented by a few populations located in some
areas of Nepal and India (Assam and western Bengal),
was documented from Early Pleistocene localities of
Java, southern China, India and Pakistan and several
Middle Pleistocene localities of the South-East Asia.
During the Late Pleistocene, R. unicornis was present
in China, Vietnam, Southern India and Sri Lanka
(Antoine, 2012). Rhinoceros sondaicus Desmarest, 1822
is currently present in limited areas of Western Java
and Southern Vietnam; but in historical times it was
widespread in several areas of South-East Asia and In-
dia. Subfossil remains of R. sondaicus were collected at
Sumatra, Borneo, Java and Malaysia, and the species
was also present in southern China (Antoine, 2012).
R. sondaicus occurred during the Pleistocene at Java,
Pandolfi
30
Malaysia, Vietnam, and Cambodia.
The genus Dicerorhinus Gloger, 1841 only occurred
in the Southeastern Asia during the Plio-Pleistocene
with extant species Dicerorhinus sumatrensis (Fischer,
1814) and the extinct species Dicerorhinus gwebinensis
Zin-Maung-Maung-Thein et al., 2008.
Some Rhinocerotina species belonging to different
lineages and biogeographic areas, i.e. C. simum (Afri-
ca), Co. antiquitatis (Northern Eurasia), R. platyrhinus
(Indian Subcontinent), acquired similar morphologi-
cal characters through their evolution, probably under
the control of similar selective pressures (Pandolfi &
Maiorino, 2016).
acKnowledgementS
I thank the reviewers and editor in chief. I also thank all
the curators of the visited museums, L. Maiorino and the
European Commission’s Research Infrastructure Action,
EU-SYNTHESYS project AT-TAF-2550, DE-TAF-3049,
GB-TAF-2825, HU-TAF-3593, ES-TAF-2997. Part of this
research received support from the SYNTHESYS Project
http://www.synthesys.info/ which is financed by European
Community Research Infrastructure Action under the FP7
“Capacities” Program.
referenceS
Antoine P.-O. (2002). Phylogénie et évolution des Ela-
smotheriina (Mammalia, Rhinocerotidae). Mémoires du
Muséum national d’Histoire naturelle, 188: 1-359.
Antoine P.-O. (2012). Pleistocene and holocene rhinocero-
tids (Mammalia, Perissodactyla) from the Indochinese
Peninsula. Comptes Rendus Palevol, 11 (2–3): 159-168.
Antoine P.-O. & Saraç G. (2005). Rhinocerotidae (Mamma-
lia, Perissodactyla) from the late Miocene of Akkaşd-
aği, Turkey. In Sen S. (ed.), Geology, mammals and
environments at Akkaşdaği, late Miocene of Central
Anatolia. Geodiversitas, 27 (4): 601-632.
Antoine P.-O., Duranthon F. & Welcomme J.L. (2003). Ali-
cornops (Mammalia, Rhinocerotidae) dans le Miocène
supérieur des collines Bugti (Balouchistan, Pakistan):
implications phylogénétiques. Geodiversitas, 25 (3): 575-
603.
Antoine P.-O., Métais G., Orliac M., Crochet J.Y., Flynn L.
J., Marivaux L., Rajpar A. R., Roohi G. & Welcomme
J. L. (2013). Mammalian Neogene biostratigraphy of
the Sulaiman Province, Pakistan. In Wang X., Flynn L.
J. & Fortelius M. (eds), Fossil Mammals of Asia: Neo-
gene Biostratigraphy and Chronology. Columbia Uni-
versity Press, New York: 400-422.
Arambourg C. (1959). Vertébrés continentaux da Miocène
supérieur de l’Afrique du Nord. Service de la Carte géolog-
ique de l’Algérie, Mémoire, 4 : 5-159.
Arambourg C. (1970). Les Vertébrés du Pléistocène de l’A-
frique du Nord. Archives du Muséum d’Histoire Naturelle,
Paris, (7) 10: 1-126.
Belyaeva E. I. (1966). Semeystvo Rhinocerotidae Owen
1845. In Vangengeym E. A., Belyaeva E. I., Garutt V.
E., Dmitrieva E. L. & Zazhigin V. S. (eds), Mlekopi-
tayushchie Eopleystozena Zapadnogo Zabaykal’ya.
Trudy Geologicheskogo Instituta Akademii Nauk
SSSR,, Moskva, 152: 92-143, 156-162.
Cerdeño E. (1992). Spanish Neogene rhinoceroses. Paleonto-
logy, 35 (2): 297-308.
Cerdeño E. (1996). Lartetotherium (Rhinocerotidae) en la
fauna con Hispanotherium del Mioceno Medio de la Re-
tama, Cuenca, España. Revista Española de Paleontología,
11 (2): 193-197.
Chen G. F. & Wu W. Y. (1976). Miocene mammalian fossils
of Jiulongkou, Cixian district, Hubei. Vertebrata PalA-
siatica, 14: 8-10.
Chow B.-S. (1963). On the skull of Dicerorhinus choukoutie-
nensis (Wang) from Choukoutien locality 20. Vertebrata
PalAsiatica, 7 (1): 62-70.
Colbert E. H. (1934). A new rhinoceros from the Siwalik
beds of India. American Museum Novitates, 749: 1-13.
Colbert E. H. (1935). Siwalik mammals in the American
Museum of Natural History. Transactions of the Ameri-
can Philosophical Society NS, 26: 1-401.
Colbert E. H. & Hooijer D. A. (1953). Pleistocene mammals
from the limestone fissures of Szechwan, China. Bulle-
tin of the American Museum of Natural History, 102: 1-134.
Deng T. (2002). The earliest known woolly rhino discovered
in the Linxia Basin, Gansu Province, China. Geological
Bulletin of China, 21 (10): 604-608.
Deng T. & Qiu Z. X. (2007). First discovery of Diceros (Pe-
rissodactyla, Rhinocerotidae) in China. Vertebrata Pala-
siatica, 45 (4): 287-306.
Deng T., Wang X., Fortelius M., Li Q., Wang Y., Tseng Z.
J., Takeuchi G. T., Saylor J. E., Saila L. K. & Xie G.
(2011). Out of Tibet: Pliocene Woolly Rhino suggests
high-plateau origin of Ice Age Megaherbivores. Science,
333: 1285-1288.
Falconer H. & Cautley P. T. (1846). Fauna antiqua sivalen-
sis, being the fossil zoology of the Siwalik Hills, in the
North of India Illustrations, part VIII: Suidae and Rhi-
nocerotidae. Smith, Elder and Co, London: pls. 69-80.
Geraads D. (1988). Révision des Rhinocerotidae (Mamma-
lia) du Turolien de Pikermi: Comparaison avec les for-
mes voisines. Annales de Paléontologie, 74: 13-41.
Geraads D. (2005). Pliocene Rhinocerotidae (Mammalia)
from Hadar and Dikika (lower Awash, Ethiopia), and a
revision of the origin of modern African rhinos. Journal
of Vertebrate Paleontology, 25 (2): 451-461.
Geraads D. (2010). Rhinocerotidae. In Werdelin L. & San-
ders W. J. (eds), Cenozoic mammals of Africa. Univer-
sity of California Press, Berkeley: 669-683.
Giaourtsakis I. X. (2009). The Late Miocene mammal fau-
na of the Mytilinii Basin, Samos Island, Greece: New
Collection 9. Rhinocerotidae. Beiträge zur Paläontologie,
31: 157-187.
Giaourtsakis I. X., Theodorou G., Roussiakis S., Athanas-
siou A. & Iliopoulos G. (2006). Late Miocene horned
rhinoceroses (Rhinocerotinae, Mammalia) from Keras-
sia (Euboea, Greece). Neues Jahrbuch für Geologie und
Paläontologie, Abhandlungen, 239 (3): 367-398.
Giaourtsakis I. X., Pehlevan C. & Haile-Selassie Y. (2009).
Rhinocerotidae. In Haile-Selassie Y. & Wolde-Gabriel
G. (eds), Ardipithecus kadabba: Late Miocene Evidence
from the Middle Awash, Ethiopia. University Califor-
nia Press, Oakland: 429-468.
Guérin C. (1966). Diceros douariensis nov. sp., un Rhinocéros
du Mio–Pliocène de Tunisie du Nord. Documents des La-
boratoires de Géologie de la Faculté des Sciences de Lyon, 16:
1-50.
Evolutionary history of Rhinocerotina 31
Guérin C. (1972). Une nouvelle espèce de Rhinocéros
(Mammalia, Perissodactyla) à Vialette (Haute-Loire,
France) et dans d’autres gisements du Villafranchien
Inférieur Européen: Dicerorhinus jeanvireti n. sp. . Docu-
ments des Laboratoires de Géologie de la Facultè des Sciences
de Lyon, 49: 53-161.
Guérin C. (1980). Les rhinocéros (Mammalia, Perisso-
dactyla) du Miocène terminal au Pleistocène supérieur
en Europe occidentale: comparaison avec les espèces
actuelles. Documents du Laboratoire de Géologie de la Facul-
té des Sciences de Lyon, 79: 1-1182.
Guérin C. (1987). Fossil Rhinocerotidae (Mammalia, Pe-
rissodactyla) from Laetoli. In Leakey M.D. & Harris J.
M. (eds), Laetoli: a Pliocene site in northern Tanzania.
Clarendon Press, Oxford: 320-438.
Guérin C. (2011). Les Rhinocerotidae (Mammalia, Perisso-
dactyla) Miocènes et Pliocènes des Tugen Hills (Kénya).
Estudios Geológicos, 67 (2): 333-362.
Handa N. & Pandolfi L. (2016). Reassessment of the middle
Pleistocene Japanese rhinoceroses (Mammalia, Rhino-
cerotidae) and paleobiogeographic implications. Paleon-
tological Research, 20 (3): 247-260.
Heissig K. (1972). Paläontologische und geologische Un-
tersuchungen im Tertiär von Pakistan. 5. Rhinocero-
tidae (Mamm.) aus den unteren und mittleren Siwa-
lik-Schichten. Bayerischen Akademie der Wissenschaften
Mathematisch-Naturwisssenschaftliche Klasse Abhandlungen
Neue Folge, 152: 1–112.
Heissig K. (1981). Probleme bei der cladistichen Analyse
einer Gruppe mit wenigen eindeutigen Apomorphien
– Rhinocerotidae. Paläontologische Zeitschriften, 55 (1):
117-123.
Heissig K. (1999). Family Rhinocerotidae. In Rössner G. E.
& Heissig K. (eds), The Miocene Land Mammals of
Europe. Pfeil, Munich: 175-188.
Heissig K. (2012). Les Rhinocerotidae (Perissodactyla) de
Sansan. In Peigné S. & Sen S., (eds), Mammifères de
Sansan. Mémoires du Muséum national d’Histoire naturelle
Paris, 203: 317-485.
Hooijer D. A. (1966). Fossil mammals of Africa no. 21.
Miocene rhinoceroses of East Africa. Bulletin of the Bri-
tish Museum(Natural History), Geology, 13 (2): 119-190.
Hooijer D. A. (1968). A rhinoceros from the Late Miocene
of Fort Ternan, Kenya. Zoologische Mededelingen, 43 (6):
77-92.
Hooijer D. A. & Patterson B. (1972). Rhinoceroses from the
Pliocene of Northwestern Kenya. Bulletin of the Museum
of Comparative Zoology, Harvard University, 144 (1):
1-26.
Kahlke R. D. & Lacombat F. (2008). The earliest immigra-
tion of woolly rhinoceros (Coelodonta tologoijensis, Rhi-
nocerotidae, Mammalia) into Europe and its adaptive
evolution in Palaearctic cold stage mammal faunas.
Quaternary Science Reviews, 27: 1951-1961.
Khan A. M., Cerdeño E., Akhtar M., Khan M. A., Iqbal A.
& Mubashir M. (2014). New fossils of Gaindatherium
(Rhinocerotidae, Mammalia) from the Middle Miocene
of Pakistan. Turkish Journal of Earth Sciences, 23: 452-
461.
Khan E. (1971). Punjabitherium, gen. nov., an extinct rhino-
cerotid of the Siwaliks, Punjab, India. Proceedings of the
Indian National Science Academy, 37 (2) A: 105-109.
Matthew W. D. & Granger W. (1923). New fossil mammals
from the Pliocene of Sze-Chuan, China. Bulletin of the
American Museum of Natural History, 48: 563-598.
Pandolfi L. & Maiorino L. (2016). Reassessment of the
largest Pleistocene rhinocerotine Rhinoceros platyrhi-
nus (Mammalia, Rhinocerotidae) from the Upper
Siwaliks (Siwalik Hills, India). Journal of Vertebra-
te Paleontology, 36 (2): e1071266 (12 pages), DOI:
10.1080/02724634.2015.1071266.
Pandolfi L., Gasparik M. & Piras P. (2015). Earliest occur-
rence of “Dihoplus” megarhinus (Mammalia, Rhinoce-
rotidae) in Europe (Late Miocene, Pannonian Basin,
Hungary): Palaeobiogeographical and biochronological
implications. Annales de Paléontologie, 101 (4): 325-339.
Tong H. (2012). Evolution of the non-Coelodonta dicerorhine
lineage in China. Comptes Rendus Palevol, 11 (8): 555-
562.
Zin-Maung-Maung-Thein, Takai M., Tsubamoto T.,
Thaung-Htike T., Egi N. & Maung-Maung (2008). A
new species of Dicerorhinus (Rhinocerotidae) from the
Plio-Pleistocene of Myanmar. Palaeontology, 51 (6):
1419-1433.
Manuscript received 14 July 2018
Received after revision 28 September 2018
Accepted 1 October 2018
Pandolfi
32
Fossilia, Volume 2018: 33-35
An
Inticetus
-like (Cetacea, Odontoceti) cheek
tooth from the Pietra Leccese (Miocene, Southern
Italy)
Emanuele Peri1, Alberto Collareta1, Gianni Insacco2 & Giovanni Bianucci1
1 Dipartimento di Scienze della Terra, Università di Pisa, via Santa Maria 53, 56126 Pisa (Italy); manuilrosso@alice.it; alberto.collare-
ta@unipi.it; giovanni.bianucci@unipi.it.
2 Museo Civico di Storia Naturale di Comiso, Comiso, 97013, Ragusa (Italy); g.insacco@comune.comiso.rg.it
introduction
Teeth in mammals are often important for the sy-
stematic study of the fossil components of this class,
like the order Rodentia, for which teeth are highly
diagnostic and many fossil species are known only
by their dental remains. However, teeth have not the
same taxonomic value in all groups of mammals: for
cetaceans, teeth often cannot define a particular spe-
cies, although some odontological features contribute
to identify certain families. Here we report the find of
an isolated cetacean tooth collected near the village
of Melpignano (Lecce Province, southeastern Italy)
from the “Pietra leccese” formation. Pietra leccese is a
Miocene (upper Burdigalian to lower Messinian) cal-
carenite that is renowned worldwide for its abundant
content of fossil marine vertebrates, which includes
turtles, bony and cartilaginous fishes, and particularly
cetaceans (both odontocetes and mysticetes). Cetace-
an remains include the holotypes of Archaeschrichtius
ruggeroi Bisconti & Varola, 2006 (Mysticeti, Eschrichti-
idae), Messapicetus longirostris Bianucci et al., 1992
(Odontoceti, Ziphiidae), Rudicetus squalodontoides
(Capellini, 1878) (Odontoceti, Kentriodontidae), and
Zygophyseter varolai Bianucci & Landini, 2006 (Odon-
toceti, Physeteroidea), as well as several odontocete
remains belonging to Squalodontidae and Eurinodel-
phinidae (Bianucci & Varola, 2014).
materialS and methodS
The cetacean dental specimen here described is sto-
red at the Museo Civico di Storia Naturale of Comiso
with catalogue number MCSNC 4457 (Insacco, 2014).
It exhibits a roughly half-circular and laterally com-
pressed crown with an almost smooth enamel surfa-
ce. This tooth bears several large accessory denticles:
along the anterior side there are four denticles, whe-
reas six denticles take place on the posterior side. The
denticles are radially positioned and slightly bowed
toward the main cusp, forming an arc. Their size gra-
dually decreases moving to the base of the crown;
moreover, in this region, there are weak subvertical
grooves on the enamel. Although most of the root is
missing, we thought that the tooth was originally dou-
ble rooted because of the presence of a slight incision
under the base of the crown. Considering the Miocene
age of MCSNC 4457, we hypothesize that it represen-
ts a cheek tooth belonging to a basal member of the
clade Neoceti (i.e. Mysticeti + Odontoceti).
In order to identify the bearer of this tooth, we com-
pared MCSNC 4457 to several groups of early bran-
ching fossil neocetes. The selection of the taxa used in
this study was mainly based on morphological simi-
larities with the Melpignano specimen. Among basal
mysticetes, Coronodon havensteini Geisler et al., 2017,
a fossil whale recently discovered from the Oligoce-
ne of the South Carolina (Ashley Formation), shows
important morphological affinities with MSNC 4457,
including the presence of several broad-based acces-
sory denticles that form an arc and smooth surface of
the enamel. We also compared MCSNC 4457 with
Squalodontidae, a family of basal odontocetes cha-
racterized by having laterally compressed postcanine
teeth with several accessory denticles (Dal Piaz, 1916;
Rothausen, 1958; 1961; 1967; 1968). Finally, we com-
pared the tooth from Melpignano with Inticetus vertizi
Lambert et al., 2018, a highly autapomorphic hetero-
dont toothed whale from the early Miocene of Peru
Bullet-pointS aBStract
• We report the find of an Inticetus-like cheek tooth from the Miocene of southern
Italy.
• We identify this specimen as belonging to cf. Inticetus sp.
• This finding brings evidence of faunal exchanges between the Mediterranean
Sea and the Pacific Ocean.
• This faunal exchange likely happened via the Central America Seaway.
KeywordS:
Inticetidae;
Mediterranean sea; palaeo-
biogeography;
Pietra leccese;
toothed whales.
Corresponding author email: manuilrosso@alice.it
How to cite: Peri et al. (2018). An Inticetus-like (Cetacea, Odontoceti) cheek tooth from the Pietra Leccese (Miocene,
Southern Italy). Fossilia, Volume 2018: 33-35. https://doi.org/10.32774/FosRepPal.20.1810.113335
Fossilia - Reports in Palaeontology
Peri et al.
34
(Chilcatay Formation). This species, that form the mo-
nospecific family Inticetidae, bears laterally flattened
cheek teeth with multiple large and bowed denticles,
that give a half-circular shape to the postcanine teeth.
reSultS and diScuSSionS
The only known mysticete with a cheek tooth mor-
phology similar to MCSNC 4457 is Coronodon haven-
steini (Geisler et al., 2017). However, accessory denti-
cles in C. havensteini are erect while those of MCSNC
4457 are weakly bowed. Furthermore, C. havensteini
has been dated to the lower Oligocene (Rupelian)
(Geisler et al., 2017) whereas the Pietra leccese lower
portions are late Burdigalian in age. Hence, MCSNC
4457 is likely too young to be considered a cheek tooth
of C. havensteini. Therefore, our morphological obser-
vations, coupled with the chronostratigraphic consi-
derations, prevent any tentative attribution of MSNC
4457 to C. havensteini or other Coronodon-like toothed
mysticetes. Alternatively, we hypothesised that MC-
SNC 4457 could belong to a member of the basal
odontocetes family Squalodontidae (e.g. Squalodon
Grateloup, 1840 and Eosqualodon Rothausen, 1968).
However, squalodontids generally have more vertical
and less pronounced denticles than our specimen, in
addition to strong ornamentation of the enamel and a
more triangular and pointed shape of the cheek teeth.
The squalodontid-like Neosqualodon Dal Piaz, 1904
exhibits more evident accessory denticles, but these are
less numerous (normally numbering three on the distal
side and two on the mesial side) and less bowed than
MCSNC 4457. Moreover, the teeth of Neosqualodon
are much smaller than MCSNC 4457. Consequently,
the morphological distance precludes the belonging of
the tooth from the Pietra leccese to any member of
Squalodontidae. The greatest similarities can be found
with the cheek teeth of Inticetus vertizi from Peru. Inde-
ed, MCSNC 4457 shares with the double-rooted cheek
teeth of Inticetus a laterally compressed and half-circu-
lar crown, with several large and bowed denticles, and
reduced ornamentation. Furthermore, in both the ho-
lotype of I.vertizi and the specimen from Pietra leccese,
slight subvertical grooves are present on the enamel.
The only morphological difference between MCSNC
4457 and the cheek teeth of I. vertizi is the smaller
number of denticles in the postcanine teeth of the lat-
ter (Lambert et al., 2018). I. vertizi is dated to the early
Miocene (Burdigalian) (Lambert et al., 2017), a time
span that is compatible with the stratigraphic range of
Pietra leccese. Therefore, based on morphological and
chronostratigraphic considerations, we identified MC-
SNC 4457 as belonging to cf. Inticetus sp.
concluding remarKS
From a palaeobiogeographical point of view, the pre-
sence of forms close to Inticetus Lambert et al., 2018
in both the southeastern Pacific Ocean and the Medi-
terranean Sea could be due to a faunistic interchange
between the Indo-Pacific Ocean and the Mediterrane-
an Sea through the Tethyan Seaway, the latter passa-
ge being still active in early Miocene times (Reuter et
al., 2009). More probably, inticetids dispersed via the
Central America Seaway: indeed, the Panama Isth-
mus did not exist until latest Miocene times (Jacobs
et al., 2004), so it is plausible that inticetids dispersed
through the Atlantic Ocean, thus recalling what has
been proposed by Bianucci et al. (2016) for the beaked
whale genus Messapicetus in late Miocene times.
In conclusion, the find of MCSNC4457 provides
new hints about the biogeographical relationships
between the Pacific and the Atlantic-Mediterranean
realms in Miocene times and suggests that our know-
ledge on the distribution patterns of the Inticetus-like
heterodont odontocetes is still fragmentary.
acKnowledgementS
Thanks to Mariano Serafini for his fundamental contribu-
tion in donating to the Museo Civico di Storia Naturale of
Comiso the cetacean tooth MCSNC 4457.We are also gra-
teful to Raymond L. Bernor and an anonymous reviewer for
their useful suggestions and constructive criticisms.
referenceS
Bianucci G. & Landini W. (2006). Killer sperm whale: a new
basal physeteroid (Mammalia, Cetacea) from the Late
Miocene of Italy. Zoological Journal of the Linnean Society,
148 (1): 103–131.
Bianucci G. & Varola A. (2014). I cetacei fossili della Pietra
leccese nei musei del Salento. Museologia Scientifica Me-
morie, 13: 130-134.
Bianucci G., Collareta A., Post K., Varola A. & Lambert O.
(2016). A new record of Messapicetus from the Pietra
leccese (late Miocene, southern Italy): antitropical di-
stribution in a fossil beaked whale (Cetacea, Ziphiidae).
Rivista Italiana di Paleontologia e Stratigrafia, 122 (1): 63-
74.
Bianucci G., Landini W. & Varola A. (1992). Messapicetus
longirostris, a new genus and species of Ziphiidae (Ce-
tacea) from the Late Miocene of “Pietra leccese” (Apu-
lia, Italy). Bollettino della Società Paleontologica Italiana,
31 (2): 261-264.
Bisconti M. & Varola A. (2006). The oldest eschrichti-
id mysticete and a new morphological diagnosis of
Eschrichtiidae (gray whales). Rivista Italiana di Paleonto-
logia e Stratigrafia, 112 (3): 447-457.
Capellini G. (1878). Della Pietra Leccese e di alcuni suoi fos-
sili. Memorie della Reale Accademia di Scienze dell’Istituto
di Bologna, 9 (3): 227-258.
Dal Piaz G. (1904). Neosqualodon nuovo genere della fami-
glia degli squalodontidi. Mémoires de la Société Paléontol-
ogique Suisse, 31:1-19.
Dal Piaz G. (1916). Gli odontoceti del Miocene del Bellu-
nese. Memorie dell’Istituto Geologico della Regia Università
di Padova, 4: 1-94.
Geisler J. H., Boessenecker R.W., Brown M. & Beatty B.L.
(2017). The Origin of Filter Feeding in Whales. Current
Biology, 27 (13): 2036-2042.
Grateloup J. P. S. (1840). Description d’un fragment de ma-
An
Inticetus
-like cheek tooth from the Pietra leccese 35
choire fossile, d’un genre nouveau de reptile (saurien).
Actes de l’Académie Nationale des Sciences Belles-Lettres et
Arts de Bordeaux, 2:201-210.
Insacco G. (2014). I Cetacei fossili del Museo Civico di Sto-
ria Naturale di Comiso (Ragusa). Museologia Scientifica
Memorie, 13: 130-134.
Jacobs D. K., Haney T. A. & Louie K. D. (2004). Genes, di-
versity, and geological process on the Pacific coast. An-
nual Review of Earth and Planetary Sciences, 32: 601-652.
Lambert O., de Muizon C., Malinverno E., Di Celma C.,
Urbina M. & Bianucci G. (2018). A new odontocete
(toothed cetacean) from the Early Miocene of Peru
expands the morphological disparity of extinct hete-
rodont dolphins. Journal of Systematic Palaeontology, 16
(12): 981-1016.
Reuter M., Piller W. E., Harzhauser M., Mandic O., Berning
B., Rögl F., Kroh A., Aubry M.-P., Wielandt-Schuster
U. & Hamedani A. (1999). The Oligo-/Miocene Qom
Formation (Iran): evidence for an early Burdigalian
restriction of the Tethyan Seaway and closure of its
Iranian gateways. International Journal of Earth Sciences
(Geol Rundsch), 98 (3): 627-650.
Rothausen K. (1958). Marine Vertebraten (Odontaspidae,
Lamnidae, Sparidae, Dermochelyidae, Squalodonti-
dae) im oberoligozänen Meeressand von Süchteln und
Düsseldorf. Fortschritte in der Geologie von Rheinland und
Westfalen, 1: 363-384.
Rothausen K. (1961). Über Microcetus, einen kleinen Squa-
lodontiden aus dem Oberoligozän. Neues Jahrbuch für
Geologie und Paläontologie, 112 (1): 106-118.
Rothausen K. (1967). Die Klimabindung der Squalodontoi-
dea (Odontoceti, Mamm.) und anderer mariner Verte-
brata. Sonderveröffentlichung des Geologischen Institut der
Universität zu Köln, 13: 157-166.
Rothausen K. (1968). Die Systematische Stellung der Eu-
ropäischen Squalodontidae (Odontoceti, Mamm.).
Paläontologische Zeitschrift, 42 (1-2): 83-104.
Manuscript received 13 July 2018
Received after revision 1 October 2018
Accepted 3 October 2018
Fossilia, Volume 2018: 37-39
First record of Monodontidae (Cetacea,
Odontoceti) in the Mediterranean Basin from the
Pliocene sands of Arcille (Grosseto, Tuscany, Italy)
Fabio Pesci, Alberto Collareta , Chiara Tinelli & Giovanni Bianucci
Dipartimento di Scienze della Terra, Università di Pisa, via Santa Maria 53, 56126 Pisa (Italy); fabio.pesci@alice.it
introduction
Extant monodontids (Cetacea: Odontoceti: Mono-
dontidae) are cold-water toothed whales that inhabit
the Artic, the North Atlantic, and the North Pacific
Oceans. The family includes two extant species: Mo-
nodon monoceros Linnaeus, 1758 and Delphinapterus leu-
cas (Pallas, 1776), commonly known as narwhal and
beluga, respectively. The fossil history of Monodonti-
dae is still scanty. Indeed, only two extinct species of
monodontids are known to date, namely Denebola bra-
chycephala Barnes, 1984 from the upper Miocene Al-
mejas Formation (Isla Cedros, Mexico), and Bohaskaia
monodontoides Vélez-Juarbe & Pyenson, 2012 from the
lower Pliocene Yorktown Formation (Virginia, USA).
In 2013, an odontocete neurocranium was recovered
from the lower Pliocene sandstones exposed in the
Arcille succession (Grosseto Province, Tuscany, Italy)
(Fig. 1). Thanks to the abundance of recent findings of
marine vertebrate skeletons, and owing to their remar-
kable state of preservation, this site is now regarded as
a genuine Pliocene marine Fossil-Lagerstätte (Tinelli et
al., 2012; Tinelli, 2013).
materialS and methodS
The odontocete partial skull from Arcille is currently
stored at the Museo di Storia Naturale dell’Universi-
tà di Pisa (MSNUP) with accession number I17602.
The list of the other odontocete skulls observed for
the current study include 7 Delphinapterus leucas (MA-
CUP AC3674; MLSUF 868; MSNUP C276; MSNUP
C277; MZUN Z315; MZUN Z324; MZUN Z1013);
8 Monodon monoceros (MACUB A4216; MACUP
AC3151; MACUP AC3488; MCSNT 773; MLSUF
870; MSNB 1387; MSNUP C274; MSNUP C275).
Institutional abbreviations
MACUB, Museo di Anatomia Comparata dell’U-
niversità di Bologna; MACUP, Museo di Anatomia
Comparata dell’Università di Pavia; MCSNT, Museo
Civico di Storia Naturale di Trieste; MZUF, Museo
“La Specola” Università di Firenze; MSNB, Museo di
Scienze Naturali di Bergamo “E. Caffini”; MSNUP,
Museo di Storia Naturale dell’Università di Pisa;
MZUN, Museo zoologico dell’Università degli Studi
di Napoli “Federico II”.
diScuSSionS and concluSionS
MSNUP I17602 is referred to Monodontidae owing
the presence of a medial exposure of the maxillae
anterior and lateral to the external bony nares [a sy-
napomorphy of monodontids according to Muizon
(1988)]. This fossil represents the first fossil occurren-
ce of the family Monodontidae in the Mediterranean
Basin. When compared with extant and extinct mo-
nodontid species, MSNUP I17602 exhibits a mosaic
of characters. It shares with Denebola brachycephala,
Delphinapterus leucas, and Monodon monoceros: an al-
most subvertical supraoccipital; a sub-circular outline
of the external bony nares. It shares with Bohaskaia
monodontoides and M. monoceros: a rounded eleva-
ted vertex; a V-shaped anterior margin of the medial
exposure of the maxillae between the premaxillae. It
shares only with D. leucas: the anteroposterior elon-
gation of the nasals (thus differing from the oval-sha-
Bullet-pointS aBStract
• MSNUP I17602 represents one of the few monodontid skull of early Pliocene
age worldwide.
• This discovery testifies the presence of a monodontid taxon in the Mediterranean
Basin.
• The fossil skull shares several characters with the skull of both extant
monodontid genera.
• Past monodontid species seems to have been adapted to subtropical climate
conditions.
KeywordS:
Monodontidae;
Pliocene;
Mediterranean Basin.
Corresponding author email: fabio.pesci@alice.it
How to cite: Pesci et al. (2018). First record of Monodontidae (Cetacea, Odontoceti) in the Mediterranean Basin from
the Pliocene sands of Arcille (Grosseto, Tuscany, Italy). Fossilia, Volume 2018: 37-39. https://doi.org/10.32774/FosRep-
Pal.20.1810.123739
Fossilia - Reports in Palaeontology
Pesci et al.
38
ped nasals of M. monoceros and B. monodontoides and
from the chevron-shaped ones of D. brachycephala); the
squamosal-parietal suture that runs at half of the hei-
ght of the temporal fossa; the transversely narrow and
almost vertical plate of the presphenoid; the posterior
end of the premaxillae taking place at about mid-len-
gth of the external bony nares (in M. monoceros the
premaxillae almost reach the nasals, whereas in B. mo-
nodontoides and D. brachycephala their posterior end is
placed far more anterior); and the vertex includes both
the nasal and frontal bones. It shares only with M. mo-
noceros: the swollen condition of the premaxillary sac
fossae; the more posterior position of the premaxillary
foramina; the arc-like disposition of the left anterior
dorsal infraorbital foramina; the well-developed crest
on the plate of the presphenoid. Moreover, a comple-
te characterization of MSNUP I17602 highlights the
presence of some autapomorphies that prevent any
possible attribution of this specimen to the four mo-
nodontid genera known to date. These characteristics
include: the depression of the premaxillae in front of
the premaxillary sac fossae; the constriction of the me-
dial exposure of the maxilla between the premaxillae
just posterior to the anterior end of the external bony
nares; the lateral expansion of the left premaxilla an-
terior to the left premaxillary foramen; the very small
size of the external bony nares; the triangular nasals
(which are even more slender and posteriorly elon-
gated than in D. leucas). Considering the subtropical
climate conditions of the Mediterranean Sea during
the early Pliocene (Bianucci et al., 2009; Prista et al.,
2015; Dominici et al., 2018), the finding of MSNUP
I17602 supports the hypothesis of an ancestral adap-
tation of Monodontidae to warm climate conditions
followed by a dispersion to high-latitude environments
which ultimately led to the origin of the modern mo-
nodontid genera.
referenceS
Barnes L. G. (1984). Fossil Odontocetes (Mammalia: Ceta-
cea) from the Almejas Formation, Isla Cedros, Mexico.
PaleoBios, 42:321-343.
Bianucci G., Sorbi S., Vaiani S. C. & Landini W. (2009).
Pliocene marine mammals from Italy: a systematic and
stratigraphic overview. International Conference on Verte-
brate Palaeobiogroprahy, Abstract book, 9-12.
Dominici S., Danise S. & Benvenuti M. (2018). Pliocene
stratigraphic paleobiology in Tuscany and the fossil
record of marine megafauna. Earth-Science Reviews,
176:277-310.
Linnaeus C. (1758) - Systema Naturae per regna tria natu-
rae, secundum Classes, Ordines, Genera, Species, cum
characteribus, differentiis, synonymis, locis. Tomus I,
10th edition: Holmiae, Laurentius Salvius, Stockholm,
Sweden, 824 p.
Muizon C. (1988). Les relations phylogénétiques des Del-
phinida (Cetacea, Mammalia). Annales de Paléontologie,
74 (4): 159-227.
Pallas P. S. (1776). Reise durch verschiedene Provinzen
des Russischen Reichs. Kayserliche Academie der Wissen-
schaften, St. Petersburg, 3:1-454.
Prista G., Agostinho R. & Cachão M. (2015). Observing the
past to better understand the future: a synthesis of the
Neogene climate in Europe and its perspectives on pre-
sent climate change. Open Geosciences, 5 (7): 65-83.
Tinelli C. (2013). Marine vertebrates from Pliocene shell
beds from Tuscany (Italy): prospecting, taphonomy, pa-
laecology and systematic palaeontology. 195pp. Unpu-
blished PhD Dissertation, Università degli studi di Pisa,
Pisa, Itay.
Tinelli C., Ribolini A., Bianucci G., Bini M. & Landini W.
Fig. 1. Measured stratigraphic sections exposed at Arcille. Modified after Tinelli (2013).
First Monodontidae in the Mediterranean Basin 39
(2012). Ground penetrating radar and palaeontology:
The detection of sirenian fossil bones under a sunflower
field in Tuscany (Italy). Comptes Rendus Palevol, 11: 445-
454.
Vélez-Juarbe J. & Pyenson N. D. (2012). Bohaskaia mono-
dontoides, a new monodontid (Cetacea, Odontoceti,
Delphinoidea) from the Pliocene of the western North
Atlantic Ocean. Journal of Vertebrate Paleontology, 32 (2):
476– 484.
Manuscript received 6 July 2018
Received after revision 28 September 2018
Accepted 2 October 2018
Fossilia, Volume 2018: 41-43
How did vertebrates sharpen their teeth? A new
perspective in bioapatite analysis
Martina Savioli1, Annalisa Ferretti1, Luca Medici2 & Daniele Malferrari1
1 Dipartimento di Scienze Chimiche e Geologiche, Università di Modena e Reggio Emilia, Via Campi 103, I-41125, Modena, Italy; martina.
savioli@unimore.it; annalisa.ferretti@unimore.it; daniele.malferrari@unimore.it
2 Istituto di Metodologie per l’Analisi Ambientale (IMAA-CNR), C. da S. Loja-Zona Industriale, I-85050, Tito Scalo, Potenza, Italy; luca.
medici@imaa.cnr.it
introduction
Analytical methods in use in crystal-chemical cha-
racterization on crystalized (natural) material can be
applied also in palaeontological investigation. X-ray
microdifraction (µ-XRD) was tested for the first time
on Ordovician conodonts from Normandy with ano-
malous overgrowth of apatite crystals on the surface
of the elements (Ferretti et al., 2017). µ-XRD allows
not only to calculate bioapatite lattice cell parameters,
but also to check for the presence of crystal preferred
orientations through comparison of XRD detected
signals changing sample orientation with respect to
X-ray beam.
When we extended our investigation to other orga-
nisms sharing bioapatite use (e.g., shark and bony fish
teeth, mammal and reptile bones, brachiopod shells,
etc.), significant variations of cell parameter values
came out. It is well known (Hughes & Rakovan, 2002)
that cell parameters are strictly dependent on chemi-
cal composition (i.e., isomorphic and iso- and hete-
ro-valent substitutions that may occur in various co-
ordination sites of bioapatite). Such information are
usually achieved through microprobe measurements.
Inductively coupled plasma mass spectrometry asso-
ciated with a laser ablation device (LA-ICPMS) may
be applied as well with the same purpose. Moreover,
LA-ICPMS is also able to acquire at the same time
(thus in the same sampling points) trace element me-
asurements. However, a mass spectrometer like any
measurement device requires an extremely accurate
calibration procedure.
When laser ablation is employed, the interaction
between laser and solid sample is complex and the
response is dependent on the sample matrix. For this
reason matrix-matched solid standards (frequently
referred to as “external standards”) are necessary to
calibrate laser ablation processes to the instrument re-
sponse. We thus tested several types of analytical stan-
dards, some of which certificated and on the market,
and others in-house prepared starting from ultrapure
reagents and certified analytical standards.
materialS and methodS
NIST 610, NIST 612 and NIST 614 are the most
commonly used analytical standard in LA-ICPMS.
Here we prepared “In-house” matrix matched stan-
dard to analyze samples characterized by a bioapatite
matrix (see next paragraph for further details). Me-
asurements were carried out using a Thermo Fisher
ICP-MS X Series II coupled with a New-Wave Laser
Ablation system. The ablation products are sent to the
inlet system of the spectrometer through an ultrapure
He flow at 500mL/min.
reSultS and diScuSSionS
Standards commonly used for LA-ICPMS analysis
are NIST 610, NIST 612 and NIST 614. However,
these standards share a glass matrix, material which
is significantly harder than bones or teeth. As a result,
when ablation conditions are applied, material remo-
ved from bioapatite samples is notably greater than
that ablated from calibrating standards. Results will
consequently be overestimated.
NIST Bone Ash and NIST Bone Meal represent a
good alternative. However, as designed for measure-
ments on water solution after mineralization, they are
unable to provide concentration ranges (i.e., if used
as a solid a single calibration point is detected). Bone
Ash and Bone Meal can be diluted mixing them with
a proper amount of an inert material (e.g., cellulose
powder); however this procedure will drive to have
standard under target concentration (i.e., a calibration
Bullet-pointS aBStract
• A new approach for the analysis of bioapatite.
• LA-ICP-MS was used to characterized samples and standards.
• “In-house” matrix matched standard were prepared to optimize chemical
analyses.
KeywordS:
bioapatite;
LA-ICP-MS;
Carcharias taurus tooth.
Corresponding author email: martina.savioli@unimore.it
How to cite: Savioli et al. (2018). How did vertebrates sharpen their teeth? A new perspective in bioapatite analysis. Fossi-
lia, Volume 2018: 41-43. https://doi.org/10.32774/FosRepPal.20.1810.134143
Fossilia - Reports in Palaeontology
Savioli et al.
42
curve that does not bracket samples concentrations).
Moreover, Bone Ash and Bone Meal do not match our
samples as far as the concentration of trace elements
is concerned.
Here two in-house made and easy to prepare stan-
dards are proposed employing analytical reagents
and/or certified analytical standards:
A mix between a micronized ultrapure synthetic hy-
droxyapatite with certified E-CRM 776-1 Firebrick
standard that can supply elements like Si, Al, Fe, Mg,
K and Na (all reported in fossilized bioapatites). The
advantage provided by this method is that the chemi-
cal composition of the “ingredients” is known and
certified and errors introduced during the preparation
should be negligible. The disadvantage is that E-CRM
776-1 Firebrick is a silicate and, thus, the final matrix
of the standard is not a pure hydroxyapatite.
A mix between a micronized ultrapure synthetic hy-
droxyapatite and ultrapure salts of the elements oc-
curring in our samples. The advantage of this method
is that it is possible to strictly monitor concentration
of each element in order to obtain standard concen-
trations that well bracket those of fossilized bioapa-
tite preserving almost completely the match between
samples and standard matrixes. On the other side, the
introduction of several salts significantly complicates
the preparation technique, possibly inducing more er-
rors.
After mixture preparation, the powder is homogeni-
zed in mortar and pressed in tablets (500 mg of pow-
der to prepare 12 ton pressed tablets with a 12 mm
diameter).
Following a procedure like that described in Nardelli
et al. (2016) we prepared also in-house made standards
for trace elements characterization. Proper amounts
of a certified solution for trace element determination
and micronized ultrapure synthetic hydroxyapatite
were mixed and homogenized in an agate mortar and
then dried at 60°C for 12 h. The resulting powder was
then re-homogenized in the agate mortar and pres-
sed at 12 tons into tablets of 12 mm diameter. Such
“standard tablets” at different elements concentration
depending on the amount of added certified solution
were then checked via LA-ICP-MS using ablation li-
nes to verify whether element distribution was homo-
geneous (Fig. 1).
concluSionS
LA-ICPMS analysis is a valuable technique able to
provide precise data on the chemical composition
(major and trace elements) even of small parts of fossil
and recent organisms using bioapatite. These data can
so be compared and discussed with cell parameters
obtained from µ-XRD. However, once again the strict
dependence on the matrix is stressed not only in terms
of chemical composition, but also in relative hardness.
In fact, as known, one of the main problems of this
LA-ICP-MS is to find the proper conditions of abla-
tion which, in turn, depend strictly on the samples and
the standards matrix. Matrix matched standards such
as the tested NIST BONE ASH and BONE MEAL,
as reported are designed for measurement on solution
after mineralization; therefore, when employed as so-
lid, cannot provide a proper calibration curve and be
used as reference to validate measurements on sam-
ples. Therefore, to apply this technique, even if some
Fig. 1. The curves show a comparison between P (A) and Ca (B) from different standards, pure hydroxyapatite, and a Carcha-
rias taurus tooth. Assuming that in Carcharias taurus tooth (Sample in the figure legend) the P and Ca content is that of a pure
hydroxyapatite (about 40% Ca, 18.5% P), we can observe that the signal from Bone Meal does not match those of the tooth,
whereas signals from Bone Ash and in house made standards are both comparable to that of the tooth. cps, counts per second;
ms, milliseconds.
A
B
New perspective in bioapatite analysis 43
steps were probably done, it is still mandatory to have
an adequate feedback system (i.e., another type of
chemical determination on solid, such as, for example,
X-ray fluorescence).
referenceS
Ferretti A., Malferrari D., Medici L. & Savioli M. (2017).
Diagenesis does not invent anything new: Precise re-
plication of conodont structures by secondary apati-
te. Scientific Reports, 7(1), article 1624. DOI:10.1038/
s41598-017-01694-4
Hughes J.M. & Rakovan J. (2002). The Crystal Structure of
Apatite, Ca5(PO4)3(F,OH,Cl). In Kohn M.J., Rakovan J.
& Hughes J.M. (eds.), Phosphates: Geochemical, Geo-
biological and Material Importance. Mineralogical So-
ciety of America, Washington DC: 1-12.
Nardelli M. P., Malferrari D., Ferretti A., Bartolini A., Sab-
batini A. & Negri A. (2016). Zinc incorporation in the
miliolid foraminifer Pseudotriloculina rotunda under
laboratory conditions. Marine Micropaleontology, 126:
42-49.
Manuscript received 15 July 2018
Received after revision 2 October 2018
Accepted 3 October 2018
Fossilia, Volume 2018: 45-47
Some species of the southeastern Italian
Miocene giant galericine
Deinogalerix
(Mammalia,
Eulipotyphla) revisited, with review of the genus
Andrea Savorelli1, Federico Masini2, Antonio Borrani2 & Paul Mazza1
1 Dipartimento di Scienze della Terra, Università degli Studi di Firenze 50121, Italy; andrea_savorelli@yahoo.it, paul.mazza@unifi.it
2 Department of Earth and Marine Sciences (DISTEM), University of Palermo, Via Archirafi 22, Palermo 90123, Italy; federico.masini@
unipa.it, antonio.borrani@unipa.it
introduction
A careful revision of fossil remains from the Terre Ros-
se of Gargano (Fig. 1A) stored at the Department of
Earth Sciences of Florence improved our knowledge
of the genus Deinogalerix Freudenthal, 1972. The aim
of this study is clearing the taxonomic status of the
specimens and, at the same time, dealing with several
issues connected with the evolutionary relationships
between the different species.
Freudenthal (1972) described the extremely derived
Deinogalerix koenigswaldi Freudenthal, 1972 and Butler
(1980) introduced four more species of Deinogalerix,
namely: Deinogalerix brevirostris Butler, 1980; Deinoga-
lerix intermedius Butler, 1980; Deinogalerix minor Butler,
1980; and Deinogalerix freudenthali Butler, 1980. Butler
(1980) indicated D. freudenthali, the smallest species of
the genus, as a hypothetical common ancestor of all
the other species. He also specified that two lineages
emerged from D. freudenthali, one passing through the
transitional D. intermedius and ending to the largest-si-
zed and most-advanced D. koenigswaldi; the other pas-
sing through D. minor and ending up with D. brevirostris
(which is smaller than D. koenigswaldi and coeval with
it). Decades later, numerous new finds of Deinogale-
rix led to the description of another primitive species,
Deinogalerix masinii Villier et al., 2013, but also to a
questioning of the taxonomic status of D. minor and
D. intermedius (Villier et al., 2013; Villier & Carnevale,
2013).
materialS and methodS
The studied material comes from infillings (“Terre
Rosse”) of karstic fissures sampled by the University
of Florence and labelled stratigraphically from the ol-
dest to the youngest: F15, F21c, NBS, P81D, F1, F8
and F9.
reSultS
The morpho-dimensional analysis performed in this
study permits to assign the material from the old fis-
sure F15 to D. freudenthali. The newly described spe-
cimens considerably improve our knowledge of the
species: D. freudenthali shares some primitive traits and
small size with the better known D. masinii (Villier et
al., 2013). Nonetheless, a handful of specimens from
F15, whose size and morphologic traits somewhat re-
call D. minor from fissure F9, are more difficult to assi-
gn to species and suggest the occurrence of a hitherto
undetected form; these have therefore been generically
indicated as Deinogalerix sp. 1.
P81D produced the richest amount of remains of
Deinogalerix of the collection of Florence. Alongside
a majority of specimens with typical features of D. in-
termedius, P81D also provided evidence of a species
with teeth roughly the size of those of D. intermedius
but with primitive traits reminiscent of D. freudenthali.
This further species was here called Deinogalerix sp. 2.
Fissure F1 and F9 record the occurrence of D. inter-
medius, accompanied by another smaller species; in F1
the latter is attested to only by a fragmental jawbone
still preserving p1, approximately the size of the man-
dibles of D. minor. In F9 the smaller species is repre-
sented by a fragmental muzzle still preserving many
of its teeth (Fig. 1B-E); it has teeth comparatively less
advanced than D. brevirostris and comes from a fissure
biochronologically older than those in which D. brevi-
Bullet-pointS aBStract
• Revision of Deinogalerix remains from the Terre Rosse of Gargano stored at the
University of Florence.
• The material was assigned mainly to D. freudenthali, D. intermedius and D. minor.
• The material improves our knowledge of the small and primitive species D.
freudenthali.
• The study confirms the validity of the lineages D. minor-D. brevirostris and D.
intermedius-D. koenigswaldi.
KeywordS:
Deinogalerix;
Terre Rosse;
late Miocene;
Galericini;
endemism.
Corresponding author address: andrea_savorelli@yahoo.it
How to cite: Savorelli et al. (2018). Some species of the southeastern Italian Miocene giant galericine Deinogalerix
(Mammalia, Eulipotyphla) revisited, with review of the genus. Fossilia, Volume 2018: 45-47. https://doi.org/10.32774/
FosRepPal.20.1810.144547
Fossilia - Reports in Palaeontology
Savorelli et al.
46
rostris is normally retrieved, therefore it is attributed
here to D. minor. Fissure F8 is virtually coeval with
F9; it yielded a single, slender mandible with features
compatible with D. intermedius.
Fissures NBS and F21c yielded a small number of
specimens. Fissure F21c may likely be contaminated,
as indicated in previous literature (Savorelli, 2013): the
large-sized and morphologically advanced p4 retrieved
in this deposit could actually belong to either D. inter-
medius or D. koenigswaldi and may have been reworked
from a more recent fissure filling. In contrast, the sin-
gle, large-sized and advanced p4 from NBS may be the
earliest occurrence of D. intermedius in the collection
of Florence.
diScuSSionS and concluSionS
The analysis performed here broadly confirms and
corroborates the framework depicted by Butler (1980);
at the same time it shows that the taxonomic revision
proposed by Villier et al. (2013) and Villier & Carne-
vale (2013), which considered valid species only the
primitive D. freundenthali and the most derived D. koe-
nigswaldi, is less convincing. Based on the new infor-
mation, D. freudenthali joins D. masinii as one of the
most primitive members of the genus. Nonetheless, D.
freudenthali is the closest to the hypothetical ancestor
of the other species described by Butler (1980). More-
over, in agreement with Butler (1980) and contrary to
Villier et al. (2013), the writers find that, similarly to
D. minor and D. intermedius, the divergence between D.
brevirostris and D. koenigswaldi is well apparent. Conse-
quently, the present study strengthens the validity of
D. minor and D. intermedius, as well as the reliability of
the phyletic lineages D. minor-D. brevirostris and D. in-
termedius-D. koenigswaldi. Members of these two lines
co-occur at least in the most recent Terre Rosse fissu-
res. Moreover, the new data, alongside recent literatu-
re (Savorelli et al., 2017), suggests that two different
species of Deinogalerix were constantly present in the
faunal assemblages of the Apulia Platform, already
since the earliest deposits.
The improved information also complicates the pi-
cture: the oldest fissures in fact contain primitive spe-
cies of unclear taxonomic and phylogenetic status. In-
deed, the fossil record of the genus remains imperfect
and the many gaps in it do not permit to utter a final
word on the origins of the various lines that characte-
rize its evolution. Nonetheless, the remains of Deino-
galerix preserved in the Department of Earth Sciences
of Florence provide substantial insight into the under-
standing of the most ancient species of the genus, they
bridge the gap of knowledge regarding D. minor, and
contribute to our understanding of the most ancient
representatives of D. intermedius. All this new infor-
mation greatly improves our comprehension of the
history of the genus Deinogalerix and, more in general,
of the evolutionary dynamics of insular species.
acKnowledgementS
We thank the Inspectors of the Archeological Superinten-
Fig. 1. A, location map of localities that yielded the fossils of the Apulia Platform Fauna. B-E, Deinogalerix minor Butler, 1980
from fissure F9, fragmental skull F9-014; B, dorsal view; C, palatal view; D, left lateral view; E, right lateral view.
Some species of
Deinogalerix
revisited, with review of the genus 47
dency of Abruzzo, M.A. Rossi and S. Agostini, for the casts
of the Gargano type specimens. We are particularly indebted
to M. Pavia, M. Delfino and G. Carnevale for granting us
access to the M010 and M013 specimens. PAULMAZZA-
RICATEN15 - Mazza P. Fondo Ateneo 2015 MIUR (the
Italian Ministry of Education, University and Research)
grants supported this study.
referenceS
Butler P.M. (1980). The giant erinaceid insectivore, Deinoga-
lerix Freudenthal, from the Upper Miocene of Gargano,
Italy. Scripta Geologica, 57: 1-71.
Freudenthal M. (1972). Deinogalerix koenigswaldi nov. gen.,
nov. spec., a giant insectivore from the Neogene of Italy.
Scripta Geologica, 14: 1-19.
Savorelli A. (2013). New data on the Cricetidae from the
Miocene “Terre Rosse” of Gargano (Apulia, Italy). Ge-
obios, 46 (1): 77-88.
Savorelli A., Masini F., Mazza P. P. A, Rossi M.A. & Ago-
stini S. (2017). New species of Deinogalerix (Mamma-
lia, Eulipotyphla) from the late Miocene of Scontro-
ne (Abruzzo, central Italy). Palaeontologia Electronica,
20.1.16A: 1-26.
Villier B. & Carnevale G. (2013). A new skeleton of the giant
hedgehog Deinogalerix from the Miocene of Gargano,
southern Italy. Journal of Vertebrate Paleontology, 33 (4):
902-923.
Villier B., Van Den Hoek Ostende L. W., De Vos J. & Pavia
M. (2013). New discoveries on the giant hedgehog Dei-
nogalerix from the Miocene of Gargano (Apulia, Italy).
Geobios, 46 (4): 63-75.
Manuscript received 15 July 2018
Received after revision 4 October 2018
Accepted 5 October 2018
Fossilia, Volume 2018: 49-51
Corresponding author: sc17278@bristol.ac.uk
How to cite: Conti et al. (2018). The oldest record of Lambeosaurinae in Europe: phylogenetic implications. Fossilia, Volu-
me 2018: 49-51. https://doi.org/10.32774/FosRepPal.20.1810.044951
Fossilia - Reports in Palaeontology
The oldest record of Lambeosaurinae in Europe:
phylogenetic implications
Simone Conti1, Albert Prieto-Márquez2, Albert Sellés Garcia2, Bernat Vila2, Àngel Galobart2,
Michael James Benton1
1School of Earth Sciences, University of Bristol, Life Sciences Building, 24 Tyndall Avenue, Bristol BS8 1TQ, United Kingdom. sc17278@
bristol.ac.uk
2 Institut Català de Paleontologia Miquel Crusafont, Universitat Autònoma de Barcelona, Carrer de l’Escola Industrial, 23, 08201, Sabadell,
Barcelona, Spain.
introduction
During the Late Cretaceous Europe was an archi-
pelago where the so called Ibero-Armorica landmass
(present-day Iberian Peninsula plus the Pyrenean and
Provence regions of France) was the largest island
(Fondevilla et al., 2016). The study of the faunas in
the Iberico-Armorican Island has evidenced a 10ma
dinosaur succession with a gradual faunal turnover.
In this, the pre-turnover dinosaur assemblages (mainly
titanosaurids, rhabdodontids, and nodosaurids) were
replaced by post-turnover assemblages, where lambeo-
saurine hadrosauroids featured predominantly (Vila et
al., 2016). Lambeosaurines in Europe are represented
by four taxa: Pararhabdodon isonensis Casanovas-Casa-
dellas et al., 1993, Arenysaurus ardevoli Pereda-Super-
biola et al., 2009, Blasisaurus canudoi Cruzado-Caballe-
ro et al., 2010 and Canardia garonnensis Prieto-Márquez
et al., 2013, belonging to three different tribes (Prie-
to-Márquez et al., 2013). All these taxa are repor-
ted from sites dated to late Maastrichtian from the
southern and northern Pyrenees. Here we update the
oldest record of European lambeosaurines.
materialS and methodS
The remains supporting this occurrence have been
collected from strata of the La Posa Formation crop-
ping out at the Els Nerets locality (early Maastrichtian,
Tremp Group, eastern Tremp Syncline, Catalonia).
The material consists of several vertebrae, a humerus,
an ulna, two ischia, two femora, a fibula and other
fragmentary elements representing two individuals,
one of which reached adulthood.
diScuSSionS and concluSionS
This lambeosaurine taxon shares characters with
Tsintaosaurus spinorhinus Young, 1958, from the Cam-
panian of China, while displaying notable differences
from Pararhabdodon isonensis from the uppermost Ma-
astrichtian of the southern Pyrenees. The phylogene-
tic position of the lambeosaurine from Els Nerets was
inferred using Maximum Parsimony and Bayesian
analyses, confirming that it is a member of Tsintao-
saurini.
The relatively small body size of this animal, com-
pared to other lambeosaurines, such as T. spinorhinus,
suggests the possibility of insularity affecting the life
history of the Iberian form.
This study reinforces the Eurasian distribution of
basal lineages of lambeosaurine hadrosaurids, indi-
cating a greater diversity in the Tsintaosaurini tribe
than previously realized. Further, the new fossils add
information to the complexity of the faunal turnover
that occurred in the Ibero-Armorican Island at the
Campanian-Maastrichtian boundary. The event was
triggered by multiple Asian dispersal waves, leading
to a Campanian fauna dominated by rhabdodontids
ornithopods, nodosaurid ankylosaurs and titanosaurs
sauropods, being subsequently replaced primarily by
lambeosaurines during Maastrichtian times.
referenceS
Casanovas-Cladellas M. L., Santafé-Llopis J. V. & Isidoro-L-
lorens A. (1993) Pararhabdodon isonensen. n. gen. n. sp.
(Dinosauria). Estudio morfológico, radiotomografico y
consideraciones biomecanicas. Paleontologia I Evoluciò.
2-27: 121-131.
Cruzado-Caballero P., Pereda-Superbiola X. & Ruiz-Omen-
cada J. I. (2010) Blasisaurus canudoi gen. et sp. nov., a
new lambeosaurine dinosaur (Hadrosauridae) from the
Latest Cretaceous of Arén (Huesca, Spain). Canadian
Journal of Earth Sciences, 47: 1507-1517.
Bullet-pointS aBStract
• Oldest lambeosaurine dinosaur from Europe.
• Close phylogenetic relationships among European and Asian lambeosaurine
dinosaurs.
KeywordS:
Dinosaur;
Lambeosaurinae;
Cretaceous;
Maastrichtian;
Europe.
Conti et al.
50
Fig. 1. Consensus tree of the Bayesian analysis, showing the phylogenetic relationships of Els Nerets specimens as basal Lam-
beosaurinae, close to Tsintaosaurus spinorhinus. The Bayesian analisis was conducted using the JC model, assuming equal rates
of changes, running 1’000’000 analyses starting from four random trees. Stationarity was achieved with a standard deviation of
split frequencies of 0.015. The matrix used for the analyses is an updated version of that of Prieto-Márquez & Gutarra (2016).
The oldest record of Lambeosaurinae in Europe: phylogenetic implications 51
Manuscript received 14 July 2018
Received after revision 15 October 2018
Accepted - October 2018
Fondevilla V., Dinarès-Turell J. & Oms O. (2016) The Chro-
nostratigraphic framework of the South-Pyrenean Maa-
strichtian succession reappraised: implications for basin
development and end-Cretaceous dinosaur faunal tur-
nover. Sedimentary Geology, 337: 55-68.
Pereda-Superbiola X., Canudo J. I., Cruzado-Caballero P.,
Barco J. L., López-Martínez N., Oms O. & Ruiz-O-
meñaca J. I. (2009) The last hadrosaurid dinosaurs of
Europe: a new lambeosaurine from the Uppermost Cre-
taceous of Aren (Huesca, Spain). Comptes Rendus Pale-
vol, 8: 559-572.
Prieto-Márquez A., Dalla Vecchia F. M., Gaete R. & Ga-
lobart À. (2013) Diversity, relationships, and bioge-
ography of the lambeosaurine dinosaurus from the
European Archipelago, with description of the new ara-
losaurin Canardia garonnensis. PLoS ONE, 8 (7): e69835.
Prieto-Márquez A. & Gutarra S. (2016) The ‘duck-billed’
dinosaurs of Careless Creek (Upper Cretaceous of
Montana, USA), with comments on hadrosaurid onto-
geny. Journal of Paleontology, 90: 133-146. doi:10.1017/
jpa.2016.42
Vila B., Sellés A. G. & Brusatte S. L. (2016) Diversity and
faunal changes in the latest Cretaceous dinosaur com-
munities of southwestern Europe. Cretaceous Research,
57: 552-564.
Young C. C. (1958) The dinosaurian remains of Laiyang,
Shantung. Paleontologica Sinica, New Series C 16: 53-
138.
Fossilia, Volume 2018: 53-56
Small and isolated: ecology and fragmentation of
Neanderthals
Marina Melchionna
1 Dipartimento di Scienze della Terra, dell’Ambiente e delle Risorse, Università degli Studi di Napoli Federico II, Italy; marina.melchionna@
unina.it
introduction
Neanderthals lived in Eurasia alongside anatomi-
cally modern humans (AMHs). The oldest evidence
of a Neanderthal population was found at Zuttiyeh
(Israel), with an age around 200,000 years ago, Tabun
(Mount Carmel, Israel) around 150,000 years (Grun
et al., 1991) and Altamura (Italy) at around 150,000
years (Lari et al., 2015). Neanderthals present unique
morphological characteristics that make them very
different from AMHs. They had a large nasal cavity,
reduced chin, and short limb proportions suggesting a
limited stature (Helmuth, 1998). Moreover, Neander-
thals had a wide chests and large lung volume (Fran-
ciscus & Churchill, 2002; Macias & Churchill, 2015).
For years scientists considered these features as adap-
tions to cold climates. Higham and colleagues (2014)
statistically placed the extinction of Homo neandertha-
lensis King, 1864 around 40 ka, almost in coincidence
with Heinrich Event 4 (HE4). This event consists in a
sudden and global shift towards colder temperatures
(Van Meerbeeck et al., 2009).
It has been demonstrated that Neanderthal popula-
tions experienced major demographic contractions
during the HE4 cold event in Northern Iberia and
Southern France (d’Errico & Goñi 2003; Sepulchre
et al., 2007). This evidence shows that, contrary to
the previous assumptions, the Neanderthal is not an
ice age species. There are different works that seem
to support this hypothesis (Finlayson & Giles, 2000;
Stewart, 2004; 2007; Bradtmöller et al., 2012).
The late contraction of H. neanderthalensis range to
southern Europe coincides with the spread of AMHs,
suggesting a possible instance for competitive exclu-
sion between the two (Banks et al., 2008; Mellars &
French, 2011). Negative interactions between Nean-
derthals and AMHs are often viewed as the potential
drivers of H. neanderthalensis extinction, as an alterna-
tive to climate change hypothesis, or a combination of
the two causes (Rey-Rodríguez et al., 2016).
Melchionna and colleagues (2018) used Species Di-
stribution Modelling (SDM) to quantify and compa-
re statistically the inferred climatic niches of Homo
sapiens Linnaeus, 1758 and H. neanderthalensis in We-
stern Eurasia during the last 8 ka of Neanderthals exi-
stence. The aim of that work was to evaluate the niche
evolution
and overlap in the two species, identifying their opti-
mal habitat patches and to which degree these patches
connected to each other.
materialS and methodS
As first step, we used fossil occurrence records (the
Stage Three Project archaeological database, van An-
del, 2002; the Canadian Archaeological Radiocarbon
Database, Gajewski et al., 2011; the Radiocarbon Pa-
laeolithic Europe Database, Vermeersch, 2017) and
paleoclimatic data (Singarayer & Valdes, 2010). Both
fossil and archaeological occurrences were used. Only
radiocarbon records computed by using Accelerator
Mass Spectrometry (AMS) were taken into account.
Dates were calibrated under the ‘IntCal13’ curve, by
using the R package ‘Bchron’ (Parnell, 2016). The fra-
mework was divided in three different temporal win-
dows, at 48 ka, 44 ka and 44 ka ago.
To model the potential distributions of H. sapiens
and H. neanderthalensis we used Species Distribution
Models (SDMs) These models allow to combine both
occurrences and climatic information to compute
the potential habitat of the species (Maiorano et al.,
2013). The final product of this procedure is a suita-
bility map. Suitability can be defined as a measure of
how much the habitat is suitable for a species to occur
in a given place and during a given time. The SDMs
computation was performed under the R software en-
vironment.
After that, we evaluated the degree of structural con-
Bullet-pointS aBStract
• Homo sapiens greater ecological plasticity allowed this species to react
better to climate changes
• AMHs maintained a more continuous occupation of its potential habitat
• Habitat reduction and fragmentation in Homo neanderthalensis must have
had dramatic consequences on its population size.
KeywordS:
Homo neanderthalensis;
Species distribution modelling;
fragmentation.
How to cite: Melchionna (2018). Small and isolated: ecology and fragmentation of Neanderthals. Fossilia, Volume 2018:
53-56. https://doi.org/10.32774/FosRepPal.20.1810.075356
Fossilia - Reports in Palaeontology
Melchionna
54
Fig. 1. Suitability analysis results. Response curves depict the variation of the probability of presence versus each variable. Blue
(green) curves referring to H. neanderthalensis (H. sapiens). Dotted lines represent the range interval over the 100 SDMs run per
species in order to account for dating uncertainty.
nectivity between optimal habitat patches as predicted
by SDMs for the two species separately, focussing on
different landscape metrics describing the number of
Patches, their area and degree of connection.
reSultS and diScuSSionS
The suitability analysis showed that Neanderthal sui-
tability is higher for 3 out of 4 climatic predictors (Fig.
1), meaning that H. neanderthalensis was better than
H. sapiens at his climatic optimum. This is not surpri-
sing, because Neanderthals originated in Eurasia, so
they were well adapted to this climatic condition. Re-
sponse curves of both species are highly overlapping,
suggesting close similarity between Neanderthals’ and
AMH’s potential climatic preferences. However, lo-
oking at the tails of the distribution, it can be noticed
that H. sapiens curves offset those of H. neanderthalensis
for three of four predictors, namely temperature du-
ring summer and both precipitation variables, sugge-
sting a wider tolerance to these predictors for H. sa-
piens (Fig 1).
The connectivity analysis showed an increase in oc-
cupied patches toward the present, but in Neander-
thals only, while the number of patches occupied by
H. sapiens remains stable (Fig. 2, top). At the same
time, the whole range of H. neanderthalensis decreases
through the process. The patches occupied by Nean-
derthals thus became smaller and more isolated (Fig.
2, bottom). This is true especially for the 44 and the 40
ka temporal windows.
Our findings seem to confirm the hypothesis of a re-
gional extinction model for North-Western Neander-
thal populations in the coldest (Northern) stretches of
its habitat (Hublin & Roebroeks, 2009), before the full
spread of AMHs in Europe, placed around 42 ka (Be-
nazzi et al., 2015). Benito et al. (2017) recently demon-
strated H. neanderthalensis most suitable environment
during the Eemian was the Mediterranean area, while
mountain ranges and continental plains showed low
habitat suitability. Our data strongly concur on these
findings. Genetic and demographic data also are con-
sistent with these notions. Neanderthals were found
to have had small population size and high mortality
rates (Trinkaus, 1995; Sørensen, 2011; Bocquet-Appel
& Degioanni, 2013).
It appears clear that the climate change played a fun-
damental role in Neanderthals demise. At the same
time, the presence of AMHs in Europe could have
been limited the latest Neanderthal populations as
well. However, a real process of habitat fragmentation
occurred in the H. neanderthalensis population and it
must have had dramatic consequences on its size.
concluSionS
Our findings show that H. sapiens had greater ecolo-
gical plasticity over Neanderthals, which probably al-
Small and isolated: ecology and fragmentation of Neanderthals 55
lowed this species to better react to climatic worsening
at 44 and then at 40 ka. On the contrary, Neander-
thals potential habitat appear to be very reduced and
fragmented during the last phase of their occupation.
Moreover, habitat reduction and fragmentation in H.
neanderthalensis suggest its population became unfit to
recover in the wake of climatic change.
acKnowledgementS
The author is grateful to Andrew Martindale for sharing
with us the CARD data. Anna Loy provided important insi-
ghts about the idea at the base of this study.
referenceS
Banks W. E., d’Errico F., Peterson A. T., Kageyama M.,
Sima A. & Sánchez-Goñi M. F. (2008). Neanderthal ex-
tinction by competitive exclusion. PLoS One, 3: e3972.
Benazzi S., Slon V., Talamo S., Negrino F., Peresani M., Bai-
ley S. E., Sawyer S., Panetta D., Vicino D., Starnini E.,
Mannino M. A., Salvadori P.A., Meyer M., Pääbo S.
& Hublin J.-J. (2015). The makers of the Protoaurigna-
cian and implications for Neandertal extinction. Science,
aaa2773.
Benito B. M., Svenning J.C., Kellberg-Nielsen T., Riede F.,
Gil-Romera G., Mailund T., Kjaergaard P.C., & San-
del B.S. (2017). The ecological niche and distribution
of Neanderthals during the Last Interglacial. Journal of
Biogeography. 44: 51–61.
Bocquet-Appel J. P. & Degioanni A. (2013). Neanderthal
demographic estimates. Current Anthropology, 54(S8):
S202-S213.
Bradtmöller M., Pastoors A., Weninger B. & Weniger G.-C.
(2012). The repeated replacement model – rapid clima-
te change and population dynamics in Late Pleistocene
Europe. Quaternary International 247: 38–49.
d’Errico F. & Goñi M. F. S. (2003). Neandertal extinction
and the millennial scale climatic variability of OIS 3.
Quaternary Science Reviews, 22(8-9): 769-788.
Finlayson C., Giles-Pacheco F. (2000). The southern Iberian
Peninsula in the late Pleistocene: geography, ecology,
and human occupation. In Stringer, C. B., Barton, R.
N. E., Finlayson, J. C. (eds.), Neanderthals on the Edge.
Oxbow Books, Oxford, pp. 139–159.
Franciscus R. G. & Churchill S. E. (2002). The costal ske-
leton of Shanidar 3 and a reappraisal of Neandertal
thoracic morphology. Journal of human evolution, 42(3):
303-356.
Gajewski K., Muñoz S., Peros M., Viau A., Morlan R. &
Betts M. (2011). The Canadian archaeological radio-
carbon data-base (CARD): archaeological 14C dates in
North America and their paleoenvironmental context.
Radiocarbon, 53: 371–394.
Grun R., Stringer C. B. & Schwarcz H. P. (1991). ESR da-
ting of teeth from Garrod’s Tabun cave collection. Jour-
nal of Human Evolution, 20(3): 231-248.
Helmuth H. (1998). Body height, body mass and surface
area of the Neandertals. Zeitschrift für Morphologie und
Anthropologie, 1-12.
Higham T., Douka K., Wood R., Ramsey C.B., Brock F.,
Basell L., Camps M., Arrizabalaga A., Baena J., Bar-
Fig. 2. Evolution of optimal habitats in Neanderthals and AMHs. Dots represent the linear distance in kilometers between
optimal patches pairs (i.e. above the 95th percentile of the suitability values predicted by the ensemble forecasting). Distance
increases from white to purple and is proportional to dots size. Row and column numbers refer to the individual patch ID. Blue
(green) columns in bar plots summarize the percentage of optimal patches pairs >4000 km apart for H. neanderthalensis (H.
sapiens) in the three time points. Dark grey (light grey) bars indicate range size of H. neanderthalensis (H. sapiens) in millions
square kilometres.
56
roso-Ruíz C., Bergman C., Boitard C., Boscato P., Ca-
parrós M., Conard N.J., Draily C., Froment A., Galván
B., Gambassini P., Garcia-Moreno A., Grimaldi S.,
Haesaerts P., Holt B., Iriarte-Chiapusso M.-J., Jelinek
A., Pardo J.F.J., Maíllo-Fernández J.-M., Marom A.,
Maroto J., Menéndez M., Metz L., Morin E., Moroni
A., Negrino F., Panagopoulou E., Pe-resani M., Pirson
S., la Rasilla de M., Riel-Salvatore J., Ronchitelli A.,
Santamaria D., Semal P., Slimak L., Soler J., Sol-er N.,
Villaluenga A., Pinhasi R. & Jacobi R. (2014). The ti-
ming and spatiotemporal patterning of Neanderthal
disappear-ance. Nature, 512: 306–309. doi:10.1038/na-
ture13621
Hortolà P. & Martínez-Navarro B. (2013). The Quaternary
megafaunal extinction and the fate of Neanderthals: An
integra-tive working hypothesis. Quaternary Internatio-
nal. 295, 69–72. doi:10.1016/j.quaint.2012.02.037
Hublin J.-J. & Roebroeks W. (2009). Ebb and flow or regio-
nal extinctions? On the character of Neandertal occupa-
tion of northern environments. Comptes Rendus Palevol,
8: 503–509.
Lari M., Di Vincenzo F., Borsato A., Ghirotto S., Micheli
M., Balsamo C., Collina C., De Bellis G., Frisia S., Gia-
cobini g., Gigli E., Hellstrom J.C., Lannino A., Modi
A., Pietrelli A., Pilli E., Profico A., Ramirez O., Riz-
zi E., Vai S., Venturo D., Piperno M., Lalueza-Foz C.,
Barbujani G., Caramelli D. & Gigli, E. (2015). The Ne-
anderthal in the karst: first dating, morphometric, and
paleogenetic data on the fossil skeleton from Altamura
(Italy). Journal of Human Evolution, 82: 88-94.
Linnaeus C. (1758) - Systema Naturae per regna tria natu-
rae, secundum Classes, Ordines, Genera, Species, cum
characteribus, differentiis, synonymis, locis. Tomus I,
10th edition: Holmiae, Laurentius Salvius, Stockholm,
Sweden, 824 p.
Maiorano L., Cheddadi R., Zimmermann N. E., Pellissier
L., Petitpierre B., Pottier J., Laborde H., Hurdu B. I.,
Pearman P. B., Psomas A., Singarayer J. S., Broenni-
mann O., Vittoz P., Dubuis A., Edwards M. E., Binney
H. A. & Guisan A. (2013). Building the niche through
time: using 13,000 years of data to predict the effects of
climate change on three tree species in Europe. Global
Ecology and Biogeography. 22: 302–317.
McGarigal K., Cushman S. A. & Ene E. (2012). FRAGSTA-
TS v4: spatial pattern analysis program for categorical
and continuous maps. In Computer Software Program
Produced By the Authors at the University of Massa-
chusetts, Amherst.
Melchionna M., Di Febbraro M., Carotenuto F., Rook L.,
Mondanaro A., Castiglione S., Diniz-Filho J. A. F. &
Raia P. (2018). Fragmentation of Neanderthals’ pre-ex-
tinction distribution by climate change. Palaeogeography,
Palaeoclimatology, Palaeoecology, 496: 146-154.
Mellars P. & French J. C. (2011). Tenfold population incre-
ase in western europe at the neandertal–to–modern hu-
man transition. Science, 333(6042): 623-627.
Parnell A. (2016). Bchron: radiocarbon dating, age-dep-
th modelling, relative sea level rate estimation, and
non-parametric phase modelling. In R package Version
4.1.1 (2015).
Rey-Rodríguez I., López-García J. M., Bennàsar M.,
Bañuls-Cardona S., Blain H. A., Blanco-Lapaz Á.,
Rodríguez-Álvarez X. P., de Lombera-Hermida A.,
Díaz-Rodríguez M., Ameijenda-Iglesias A., Agustí J.
& Fábregas-Valcarce R. (2016). Last Neanderthals and
first Anatomically Modern Humans in the NW Iberian
Peninsula: Climatic and environmen-tal conditions in-
ferred from the Cova Eirós small-vertebrate assemblage
during MIS 3. Quaternary Science Review, 151: 185–197.
Sepulchre P., Ramstein G., Kageyama M., Vanhaeren M.,
Krinner G., Sánchez-Goñi M.-F. & D’errico F. (2007).
H4 abrupt event and late Neanderthal presence in Ibe-
ria. Earth and Planetary Science Letters, 258: 283–292.
Singarayer J. S. & Valdes P. J. (2010). High-latitude climate
sensitivity to ice-sheet forcing over the last 120kyr. Qua-
ternary Science Review, 29: 43–55. doi:10.1016/j.quasci-
rev.2009.10.011
Sørensen B. (2011). Demography and the extinction of Eu-
ropean Neanderthals. Journal of Anthropological Archaeo-
logy, 30(1): 17-29.
Stewart J. R. (2004). Neanderthal–modern human compe-
tition? A comparison between the mammals associated
with Mid-dle and Upper Palaeolithic industries in Eu-
rope during OIS 3. International Journal of Osteoarchaeol.
14: 178–189. doi: 10.1002/oa.754
Stewart J. R. (2007). Neanderthal extinction as part of the
faunal change in Europe during Oxygen Isotope Stage
3. Acta Zoologica Cracoviensia - Series A: Vertebrata, 50A:
93–124. doi:10.3409/000000007783995372
Trinkaus E. (1995). Neanderthal mortality patterns. Journal
of Archaeological Science, 22(1): 121-142.
van Andel T. H. (2002). The Climate and Landscape of the
Middle Part of the Weichselian Glaciation in Euro-
pe: The Stage 3 Project. Quaternary Research, 57: 2–8.
doi:10.1006/qres.2001.2294
Van Meerbeeck C. J., Renssen H. & Roche D. M. (2009).
How did Marine Isotope Stage 3 and Last Glacial Ma-
ximum cli-mates differ?–perspectives from equilibrium
simulations. Climate of Past, 5: 33–51.
Vermeersch P. M. (2017). Radiocarbon Palaeolithic Europe
Database, Version 24. Available at: http://ees.kuleu-
ven.be/geography/projects/14c-palaeolithic/index.
html
Melchionna
Manuscript received 5 August 2018
Received after revision 16 October 2018
Accepted 22 October 2018
Fossilia, Volume 2018: 57-60
How to cite: Mondanaro (2018). Mechanisms of competition in mammals during the Late Neogene. Fossilia, Volume
2018: 57-60. https://doi.org/10.32774/FosRepPal.20.1810.085760
Fossilia - Reports in Palaeontology
Mechanisms of competition in mammals during
the Late Neogene
Alessandro Mondanaro
Dipartimento di Scienze della Terra, Università degli Studi di Firenze 50121, Italy; alessandro.mondanaro@unifi.it
Dipartimento di Scienze della Terra, dell’Ambiente e delle Risorse, Università degli Studi di Napoli Federico II, Italy
introduction
Megaherbivore mammals are traditionally defined as
species above 1000 kg (du Toit & Owen Smith, 1989;
Fritz et al., 2011). In modern-day ecosystems, this eco-
logical category includes few species such as rhinos,
elephants, the hippopotamus, gaur, yak, and the giraf-
fe, and it is geographically restricted to sub-Saharian
Africa and Asia. However, before the Late Pleistoce-
ne megafauna extinction, megaherbivores were much
more diverse. The so-called mammal megafauna was
then represented by mastodonts, mammoths, ground
sloths, giant armadillos, a number of notoungulates,
very large bovids, deer, and camels, plus giant kanga-
roos and other marsupials. Megaherbivores control
the consumption of primary production excluding
smaller species from accessing limited resources, and
exposing them to an increased predation risk by clea-
ring thickets of vegetation the small game usually use
to hide from predators. Moreover, in virtue of their
large body size, megaherbivores have virtually no pre-
dator (Fritz et al., 2011). Consequently, when abun-
dant, megaherbivores also cause a negative effect on
predators limiting their availability of prey. This ecolo-
gical mechanism is known as “apparent competition”
and describes the indirect competition for limiting
resources. In this work, we verified the idea that ap-
parent competition of megaherbivores on carnivores
applies to Neogene to Recent large mammals at large
geographic and temporal scales. Furthermore, we te-
sted the idea that the diversity of megaherbivores posi-
tively affected the diversity of sabertoothed cats. Such
species were deemed to have specialized to kill largest
among prey and are said to have gone extinct once
the latter vanished by the end of the Pleistocene. In
contrast, evidence coming from studies of their ena-
mel isotopic composition (Feranec, 2005), long-term
analyses of their prey consumption style (DeSantis et
al., 2012), modelling of prey selection in extinct guilds
of carnivores (Randau et al., 2013) suggested a scarce
evidence of predation mechanism by sabertooths on
megaherbivores. Taking in mind this information, we
tested for the association between sabertooths like Ho-
motherium Fabrini, 1890 and Smilodon Lund, 1842 and
megaherbivore fossil remains (Raia et al., 2007; Van
Valkenburgh et al., 2016).
materialS and methodS
We downloaded from the Paleobiology Database,
NOW Database and Pangaea Database, Neogene
fossil occurrences of mammals belonging to Artio-
dactyla, Perissodactyla, Proboscidea, Carnivora and
Creodonta. The geographical range of mammal oc-
currences covers the entire Eurasiatic region plus the
Africa continent. We supplemented these data with
occurrence records from Raia et al. (2009) and other
published sources. For each species, age estimates and
geographical paleocoordinates of individual occurren-
ces at fossil sites were recorded. Overall, the occurren-
ces dataset includes 655 artiodactyls, 114 carnivores, 4
creodonts, 163 perissodactyls, 67 proboscideans, for a
total of 1003 species spanning from the earliest Mioce-
ne to the Holocene, distributed over 3021 fossil locali-
ties. For each species, we compiled body size estimates
from different databases. Species were divided in four
ecological categories based on both diet and estima-
ted body mass: Megaherbivores (Mega), Herbivores
Bullet-pointS aBStract
• Although rare, megaherbivores negatively affect species diversity at several
trophic levels.
• In contrast, it has been suggested that they favour coexistence among top pre-
dators.
• We show megaherbivores controlled ecosystem functioning in Neogene large
mammals.
• Carnivores were a factor in controlling the diversity of small prey.
KeywordS:
Predation to prey ratio;
Megaherbivores;
Apparent competition
Mondanaro
58
(Herb), Carnivores (Carn) and Sabertooths (Sab).
Then, we divided all record in 2-million-years-long
temporal intervals maintaining a reasonably dense
record for each interval. For each species and within
each time bin, we constructed minimum convex poly-
gons (MCP, Carotenuto et al., 2010) starting from its
fossil occurrences (Fig.1). We overlaid a 500 × 500 km
grid cell resolution on each projected continent, sam-
pling regions in an equal area context.
The use of polygons overcomes problems generated by
sampling inequality per species and geographic area,
by adding cells to the species presence where no fossil
occurrence is indeed present, but still within the mini-
mal range of species geographic extent. Similarly, by
using a geographic grid all the fossil localities falling
within a given cell (i.e. within an area of 25,000 km2)
in a given time intervals are collapsed in a single fau-
nal list. In this way, the effect of unequal sampling and
taphonomic effects across fossil sites is avoided. We
performed all analyses twice, both by using the raw
fossil occurrence (henceforth “occurrence record”),
and by using MCP polygons to attribute species to
cells (henceforth “polygon record”). At this stage, for
each cell and time bin, we excluded cells with < 5 spe-
cies overall, or lacking any predator or prey, either.
After that, we computed the ratio between the num-
ber of “predators” (Carnivores plus Sabertooths) and
“preys” (Meso- and Megaherbivores) for each geo-
graphic cell within a given time bin. We also compu-
ted the body size range of Predators and Prey for each
cell within each time bin in order to retrieve the degree
of overlap between Predators and Prey body mass di-
stributions (PPR-overlap).
With these variables, we performed six different regres-
sions using the number of species in each category and
their estimated body sizes per cell, and separately per
time bin. The regression (1) of predator-to-prey ratio
(PPR) against the number of megaherbivores (Mega)
was calculated to test for the effect of apparent com-
petition of the latter on Predators. Predators (Pr) were
regressed (2) against Herbivores (Herb) to verify for
the relationship between the richness of predators and
non-megaherbivore prey. The regression (3) between
Mega and Herb was computed to test for competition
between species belonging to these categories, to test
the idea that megaherbivores did control the diversity
of Herb. The richness of Sabertooths (Sab) was regres-
sed (4) against Mega in order to test the idea that sa-
bertooths preferentially preyed upon megaherbivores.
Similarly, the diversity of large carnivores (Pr > 100)
was regressed against Mega (5). Eventually, we regres-
sed (6) PPR against Overlap in order to verify if an
increment in degree of overlap is correlated to a higher
chance of predation on megaherbivores, under the ob-
servation that larger predator might tackle down com-
paratively larger prey (Van Valkenburgh et al., 2016).
All of the six regressions were controlled for spatial
autocorrelation by using GLS models. In details, we
fitted the related empirical semivariograms with 4 mo-
dels (Gaussian, Spherical, Rational Quadratic, Expo-
nential) and then updated an OLS regression by these
4 spatial correlation structures. The outcomes of these
five models (OLS and the 4 spatially structured) were
then compared by means of ANOVA.
Consecutive intervals within the same geographical
place share a number of species. This means that the
data could be temporally autocorrelated, thereby ori-
ginating spurious associations between the variables.
To address the issue of temporal autocorrelation, we
used the autoregressive integrated moving average
(ARIMA) model. ARIMA works by regressing a va-
riable point value on previous (older) datapoints, at
some (fitted) distance (i.e. lag). The best lag between
variables was estimated via cross-correlation, and the
existence (and removal thereof) of temporal autocor-
relation was assessed by means of Breush-Godfrey
test (McMurry & Politis, 2015). To produce ARIMA
models, we used the polygons record to maximize the
Fig. 1. The geographical distribution of fossil localities included in the analyses (left). Geographical coordinates were rotated to
the present for plotting purposes. To the right, for each species it is plotted the minimum convex polygon including all species
occurrences (right).
Mechanisms of competition in mammals during the Late Neogene 59
number of datapoints. However, rather than using the
500 km wide cells we opted for 2000 × 2000 km cells
and selected only cells possessing at least 10 species
and at least one carnivore species per time bin, for at
least 6 time bins. We performed the ARIMA regres-
sions of PPR against Mega, Pr against Herb, Mega
against Herb, and Pr > 100 against Mega.
reSultS and diScuSSionS
The results were qualitatively very similar using ei-
ther the polygon and the occurrence record. The poly-
gon record is much more dense and less affected by
sampling issues, and was therefore used to perform
ARIMAs.
As far as the six regression are concerned, the preda-
tor to prey ratio (PPR) is significantly and negatively
correlated to the number of megaherbivores in one
fourth of the intervals. No positive relationship ap-
plies. The number of predators (Pr) is negatively and
significantly related to the number of mesoherbivores
(Herb) in 6 out of 8 intervals (75%) (Tab. 1). Nearly
two-thirds of the times (5 times in 8 intervals, 62.5%)
the richness of megaherbivores (Mega) is inversely and
significantly related to herbivore (Tab. 1). There is no
positive relationship between the diversity of saberto-
oths (Sab) and Mega. The richness of large predators
(species > 100 kg in body size) is not significant asso-
ciated to the number of megaherbivores (Tab. 1). The
regression between Mega and Pr > 100 is significant
and negative. Finally, the degree of overlap between
the body size distribution of predators and prey is
positively and significantly related to PPR six times
(66.7%) (Tab. 1). A significant and negative relation-
ship occurs once (Tab. 1). As regards the first (most
recent) interval, all the regression results are consistent
with the apparent competition theory. The spatial di-
stribution of PPR in the Old World during the last in-
terval shows no significant spatial autoregression. The
results of ARIMA regressions confirm the existence
of a negative relationship between the number of pre-
dators (Pr) and mesoherbivores (Herb) in four out of
five valid cells. The number of megaherbivores is ne-
gatively associated to the number of mesoherbivores
(Mega-Herb) in three cells out of four. PPR is always
negatively associated to Mega. Finally, the number of
megacarnivores (i.e. predators above 100 kg in body
mass) is negatively associated to the number of me-
gaherbivores twice, and positively associated as many
times. Cross-correlation applied to the residuals consi-
stently shows a lag of 1 to 3 intervals, and mostly of 2
(i.e. 4 Myr) between herbivores and predators and me-
gaherbivores respectively, meaning that Pr and Mega
negatively affected the diversity of mesoherbivores in
successive temporal intervals.
These results indicate that large carnivores paid a
price to the ecological dominance of megaherbivores.
As megaherbivores diversified, the biomass available
Regression
model (y-x)
Significant Non-
significant
Positive Negative
PPR-Mega 0 2 7
Pr-Herb 0 6 2
Mega-Herb 0 5 3
Sab-Mega 0 1 7
Pr > 100-Mega 0 1 7
PPR-Overlap 6 1 2
to carnivores reduced, exactly because megaherbivores
are hard to kill, and smaller herbivores were signifi-
cantly outcompeted by the larger species. Indeed, ra-
ther than the apparent competition of megaherbivores
on carnivores, the most robust generalizations we deri-
ve from this study is that the diversity of mesoherbivo-
res is negatively affected by the diversity of both carni-
vores and megaherbivores. This implies that predation
and direct competition increases extinction probability
in small herbivores, as often suggested to occur for li-
ving prey species (Fritz et al., 2002; Malhi et al., 2016).
Some sabertooths were probably able to dispatch ju-
veniles of very large prey species such as mammoths
or ground sloths. This was suggested to exercise a top-
down control on megaherbivores (Van Valkenburgh
et al., 2016). While killing juvenile megaherbivore is
perfectly feasible for a top predator, and not contradi-
cted by our data, we suggest these were probably not
their most profitable prey, and more importantly, me-
gaherbivores diversity did not sustain higher carnivore
diversity.
concluSionS
We found evidence that, in the long run, such pervasi-
ve dominance of megaherbivores translates into higher
extinction rate upon (primarily) small prey and (secon-
darily) predatory species, thereby altering predator to
prey ratios (Raia et al., 2007; Meloro & Clauss, 2012).
Such fact is further conceivable considering that small
prey populations tend to be limited by predation, while
larger species are mainly controlled by resources (Ter -
borgh et al., 2010). It must be noted that very large pre-
dators and sabertooths were much less influenced by
the diversity of megaherbivores than mesoherbivores.
Together with the quite consistent, positive relation-
ship between body size overlap and PPR, this indica-
tes that very large carnivores did actually go for larger
prey than other carnivores on average. However, this
is also consistent with their larger average body size,
and not just with any preference for megaherbivores.
Indeed, large predators do not specialize on the larger
prey, they just exploit a wider prey spectrum (Radloff
& du Toit, 2004).
Tab. 1. Summary statistics for the regressions of the number
of species within ecological categories per cell.
Mondanaro
60
acKnowledgementS
I’m grateful to Lorenzo Rook, Pasquale Raia e Franscesco
Carotenuto who shared with me the ideas ehich are the base
of this work.
referenceS
Carotenuto F., Barbera C. & Raia P. (2010). Occupancy, ran-
ge size, and phylogeny in Eurasian Pliocene to recent
large mammals. Paleobiology, 36, 399–414.
DeSantis L. R. G., Schubert B. W., Scott J. R. & Ungar
P. S. (2012). Implications of diet for the extinction of
saber-toothed cats and American lions. PLoS ONE,
7: e52453. http://dx. doi.org/10.1371/journal.
pone.0052453
du Toit J. T. & Owen-Smith N. (1989). Body size, popula-
tion metabolism, and habitat specialization among
large African herbivores. American Naturalist, 133 (5),
736–740. http://dx. doi.org/10.2307/2462079.
Feranec R. S. (2005). Growth rate and duration of growth
in the adult canine of Smilodon gracilis, and inferences
on diet through stable isotope analysis. Bulletin of the
Florida Museum of Natural History, 45, 369–377.
Fritz H., Duncan P., Gordon I. & Illius A. (2002). Megaher-
bivores influence trophic guilds structure in African
ungulate communities. Oecologia, 131, 620–625. http://
dx.doi.org/10.1007/s00442-002-0919-3.
Fritz H., Loreau M., Chamaillé Jammes S., Valeix M. &
Clobert J. (2011). A food web perspective on large her-
bivore community limitation. Ecography, 34, 196–202.
Malhi Y., Doughty C. E., Galetti M., Smith F.A., Svenning
J.-C. & Terborgh J. W. (2016). Megafauna and ecosy-
stem function from the Pleistocene to the Anthropocene.
Proceedings of the National Academy of Sciences, 113: 838–
846. http://dx.doi.org/10.1073/pnas.1502540113.
McMurry T. L. & Politis D. N. (2015). High-dimensional
autocovariance matrices and optimal linear prediction.
Electronic Journal of Statistics, 9: 753–788.
Radloff F. G. T., du Toit J. T. & Johan T. (2004). Large pre-
dators and their prey in a southern African savanna: a
predator’s size determines its prey size range. Journal of
Animal Ecology, 73: 410–423.
Raia P., Meloro C. & Barbera C. (2007). Inconstancy in
predator/prey ratios in Quaternary large mammal
communities of Italy, with an appraisal of mechani-
sms. Quaternary Research, 67: 255–263. http://dx.doi.
org/10.1016/j.yqres.2006.10.005.
Randau M., Carbone C. & Turvey S. T. (2013). Canine evo-
lution in sabretoothed carnivores: natural selection or
sexual selection? PLoS ONE, 8, e72868.
Terborgh J., Holt R. D. & Estes J. A. (2010). Trophic casca-
des: what they are, how they work, and why they mat-
ter. In Terborgh, J., Estes, J.A. (Eds.), Trophic Cascades:
Predators, Prey and the Changing Dynamics of Nature.
Island Press, USA, pp. 1–18.
Van Valkenburgh B., Hayward M. W., Ripple W. J., Meloro
C. & Roth V. L. (2016). The impact of large terrestrial
carnivores on Pleistocene ecosystems. Proceedings of the
National Academy of Sciences, 113 (4): 862–867. http://
dx.doi.org/10.1073/pnas.1502554112.
Manuscript received 5 August 2018
Received after revision 16 October 2018
Accepted 22 October 2018
The “Brunella” Project: preparation and study of a
mysticete from the Early Pliocene of Tuscany
Roberta Scotton1, Renzo Bigazzi1, Simone Casati2, Giuseppe D’Amore1, Sylvia Di Marco1,
Luca Maria Foresi3, Elizabeth Koenig4, Luca Ragaini5, Jacopo Tabolli6, Massimo Tarantini7,
Giandonato Tartarelli8 & Michelangelo Bisconti9
1 Istituto di Studi Archeo-Antropologici, Via delle Cascine 46, 50018, Scandicci (Firenze); robertascotton@yahoo.it; renzo.bigazzi@
teletu.it; g_damore@libero.it; sylvia_dimarco@libero.it
2Gruppo AVIS di Mineralogia e Paleontologia Scandicci, Piazza Vittorio Veneto 1, 50018, Scandicci (Firenze); simonecasati@alice.it
3Dipartimento di Scienze Fisiche, della Terra e dell’Ambiente, Università degli Studi di Siena, via Laterina 8, 53100, Siena; luca.foresi@
unisi.it
4Banfi s.r.l., Castello di Poggio alle Mura, 53024, Montalcino (Siena); elizabeth.koenig@banfi.it
5Dipartimento di Scienze della Terra, Università di Pisa, via Santa Maria 53, 56126, Pisa; ragaini@dst.unipi.it
6Soprintendenza Archeologia, Belle Arti e Paesaggio per le province di Siena, Grosseto, Arezzo, via di Città 138/140, 53100, Siena; jacopo.
tabolli@beniculturali.it;
7Soprintendenza Archeologia, Belle Arti e Paesaggio per la città metropolitana di Firenze e le province di Pistoia e Prato, Piazza de’ Pitti 1,
50125, Firenze; massimo.tarantini@beniculturali.it
8Scuola Normale Superiore, Piazza Cavalieri, 56100, Pisa; tartarelli@sns.it
9San Diego Natural History Museum, 1788 El Prado, CA 92101, San Diego, United States of America; michelangelobisconti@gmail.
com
introduction
In 2007, the skeleton of a mysticete was uncovered
in the Banfi’s vineyard at the locality of Poggio alle
Mura, near Montalcino (Siena). The excavation of
this whale resulted in the discovery of a partially ar-
ticulated skeleton including skull, both dentaries, part
of the vertebral column, one scapula, parts of the fo-
relimbs and ribs. Since the excavation this whale has
been referred to with the nickname “Brunella”. The
excavation revealed also mollusks, barnacles and pa-
laeobotanical remains. In addition, a preliminary in-
vestigation of the stratigraphy at the site was carried
out and sediment samples for micropalaeontological
analyses were gathered from the stratigraphic level
where the whale was found and from different layers
of the stratigraphic section. The excavation occurred
during an important enological event in Montalcino
and the discovery of the whale was presented on na-
tional television channels (RAI) and onto numerous
national and international newspapers and journals
(e.g., Anonymous, 2007). In the following years, the
fossils resulting from the 2007 excavation were stored
in a warehouse at the Banfi’s property at Poggio alle
Mura. In 2016, the Soprintendenza Archeologica del-
la Toscana decided to begin a new project focused on
the preparation, study and possible exhibition of this
whale on the basis of early reports suggesting that it
could be an important step towards shedding light on
mysticete evolution, and on palaeobiogeographic and
palaeoecological conditions at the site of the disco-
v er y.
materialS and methodS
The mysticete specimen and the sediment samples
taken at the discovery site are preserved in the labo-
ratory at Poggio alle Mura (Montalcino, Siena Pro
vince). The geological section at the site was studied
during the excavation in 2007 and samples were obtai-
Bullet-pointS aBStract
• A project is being carried out at Poggio alle Mura that will allow the
preparation and study of a balaenopterid whale from the early Pliocene of
Tuscany together with its associated biota.
• Up to now, the project resulted in the preparation of 18 vertebrae, the skull and
an ulna of the whale.
• The associated biota includes 18 mollusk species, 2 sea urchin species, 10 shark
teeth and thousands of fish remains from the sediment surrounding the skull.
• A wealth of educational activities is being carried out at the laboratory of
Poggio alle Mura that are directed to schools and citizens.
KeywordS:
mysticeti;
Pliocene;
Tuscany;
anatomy;
paleoenvironment
Fossilia, Volume 2018: 61-63
Corresponding author: robertascotton@yahoo.it
How to cite: Scotton et al. (2018). The “Brunella” Project: preparation and study of a mysticete from the Early Pliocene of
Tuscany. Fossilia, Volume 2018: 61-63. https://doi.org/10.32774/FosRepPal.20.1810.156163
Fossilia - Reports in Palaeontology
Fig. 1. The “Brunella” project. A, the skull of the mysticete in the slab where it was preserved together with abundant mollusks.
B, another view of the skull with the bones digitally colored in blue for closer individuation. C, the skull with rostrum digitally
colored in orange and neurocranium in blue. D, a group of lumbar vertebrae, caudal vertebrae and chevrons in the slab where
they are preserved. E, a student of a summer school during preparation activities on “Brunella”’s ribs.
Scotton et al.
62
ned from different layers to be used for micropalaeon-
tological analyses. Sediments were washed and sieved
to be examined under microscopy in search for micro-
fossils. The mysticete bones were subject to mecha-
nical preparation; use of alcohol and oxidized water
supplemented the mechanical preparation to facilitate
sediment removal. The sediment removed from the
bones was washed and sieved to be analyzed under a
stereomicroscope with magnification of 10x and 20x.
Standard sediment samples were 1 l in volume cor-
responding to 1476 g in weight. All the microfossils
were saved in dedicated containers. During the micro-
excavation of the bones, the chemical product used in
2007 to consolidate the skeleton was removed by ap-
plication of acetone and a new bicomponent chemical
product was then applied to definitively consolidate
the bones. Mollusk shells were mechanically polished.
The total body length of the whale was calculated by
using dedicated regression equations (Pyenson et al.,
2013) based on a preliminary estimation of the length
of the right dentary.
diScuSSionS and concluSionS
The new project involved both public and private
institutions in a joint effort with the following goals:
prepare the specimen; obtain a chronostratigraphic
assessment of its age; describe and analyze the pala-
eoenvironment at the site of the discovery based on
mollusk, foraminifer and sediment analyses; descri-
be the specimen and investigate into its systematic,
phylogenetic and palaeobiogeographic relationships;
exhibit the skeleton within an existent or new location
(e.g., a museum); and, realize a series of educational
programs around the specimen, such as a summer
school focused on the preparation of paleontological
remains open to undergraduate students. The project
was funded by the Banfi Society following the recently
established Art Bonus scheme of the MiBACT (Italian
Ministry of Cultural Heritage and Tourism). To our
knowledge, this is the first project of this kind focused
on a fossil vertebrate to be developed in Italy.
The project started following separate agreements
between the Archaeological Superintendency of Tu-
scany, the Banfi Society, and the Istituto di Studi Ar-
cheo-antropologici (ISA), resulting in two editions of
a summer school realized in 2016 and 2017 involving
teachers from three universities of Tuscany, the Scuola
Normale Superiore of the University of Pisa, the ISA,
the Museo di Storia Naturale di Milano, and the San
Diego Natural History Museum (Fig. 1). The third
edition of the summer school is currently under pre-
paration and will take place by the end of 2018. The
school has received the official endorsements by the
Manuscript received 13 July 2018
Received after revision 25 September 2018
Accepted 2 October 2018
The “Brunella” Project 63
Società Paleontologica Italiana (SPI) in previous edi-
tions, the Dipartimento di Scienze Fisiche, della Terra
e dell’Ambiente of the University of Siena, and the
Associazione Italiana per lo Studio del Quaternario
(AIQUA). The following subjects have been included
in the programs of the school: mysticete osteology,
evolution and biogeography; general biological and
geological patterns in the Mediterranean Pliocene;
Miocene and Pliocene mollusk evolution; mysticete
taphonomy; palaeogeographic and palaeoenviron-
mental evolution of Tuscany during the Pliocene;
paleontological excavation techniques; techniques
of preparation and restoration of fossil vertebrates;
techniques of photogrammetry and tridimensional
modelling of fossil vertebrates; and museology. Stu-
dents from different universities came from northern,
central and southern Italy sharing their cultural back-
grounds: Geology, Natural Sciences and Conserva-
tion of Cultural Heritage. The Banfi Society provided
the location for the classes, the warehouse for the
preparation laboratory of the fossil whale (as well as
food). RAI3 (national television channel), local new-
spapers and web journals provided media coverage on
the different activities of the school. Presently (June
2018), the preparation and restoration of the speci-
men is in progress. The technicians are using mecha-
nical preparation instruments to remove the matrix
from the bones and a diverse array of chemical pro-
ducts to consolidate and preserve the fossil. The pre-
servation is variable: the vertebrae and the posterior
portion of the skull are in good conditions while the
rostrum is badly damaged by post-mortem processes.
The scientific study of “Brunella” is still in progress.
Preliminary investigations revealed that: it belonged
to the family Balaenopteridae; it does not belong to
any of the living taxa; based on the length of its den-
tary, it was estimated that the total body length of
“Brunella” should have been between c. 6 and c. 7.5
m and its weight was c. 5000 kg; the age of this wha-
le was preliminarily assessed based on foraminifers
and mollusks and was constrained between 3.7 and
4.5 Ma. “Brunella” was framed in publications on the
taphonomy and stratigraphic palaeobiology of baleen
whales from Italy (Dominici et al., 2009; 2018). Gi-
ven its degree of completeness and preservation, it is
expected that “Brunella” will shed further light on the
evolution of balaenopterid mysticetes from the Early
Pliocene of the Mediterranean basin.
acKnowledgementS
The “Brunella” project was made possible by the interest
and action of several individuals: SC discovered the spe-
cimen together with Franco Gasparri; Menotti Mazzini
(former technician of the University of Florence) directed
the 2007 excavation; Enrico Viglierchio, Remo Grassi (all
at Banfi s.r.l.), Giorgio Carnevale (University of Turin),
Cristiano Dal Sasso (Natural History Museum of Milan)
provided valuable insights and help during the research and
the field school editions.
referenceS
Anonymous (2007). The whales of Italy. Science, 316: 179.
Dominici S., Cioppi E., Danise S., Betocchi U., Gallai G.,
Tangocci F., Valleri G. & Monechi S. (2009). Mediter-
ranean fossil whale falls and the adaptation of mollu-
sks to extreme habitats. Geology, 37: 815–818.
Dominici S., Danise S. & Benvenuti M. (2018). Pliocene
stratigraphic paleobiology in Tuscany and the fossil re-
cord of marine megafauna. Earth Sciences Reviews, 176:
277-310.
Pyenson J. D., Goldbogen J. A., Shadwick R. E. (2013).
Mandible allometry in extant and fossil Balaenopteri-
dae (Cetacea: Mammalia); the largest vertebrate skele-
tal element and its role in rorqual lunge feeding. Biologi-
cal Journal of the Linnean Society, 108: 586-599.
