Three islands, three worlds: Paleogeography and evolution of the vertebrate fauna from the Balearic Islands

Abstract

The Balearic Islands are an archipelago located in the Western Mediterranean Sea. Their isolation from the mainland allowed the establishment of different faunas on each island. In Mallorca, the Pliocene fauna was composed of the so-called Myotragus-fauna (mainly consisting of a bovid, a glirid and a soricid). In Menorca, it was constituted by the giant rabbit-fauna (mainly consisting of a giant rabbit and a tortoise), and in the Pityusics by a tortoise, a lizard and two rodents. A main faunal turnover took place during the Late Pliocene or Early Pleistocene: the Myotragus-fauna reached Menorca and replaced the giant rabbit fauna. In the Pityusics, all mammals and the tortoise became extinct before the Late Pleistocene for unknown reasons, leaving birds and the lizard as the only vertebrates of these islands. Almost all the endemic vertebrates of the Balearics became extinct probably due to the first human arrival to the islands.

Full text

Quaternary International 182 (2008) 135–144
Three islands, three worlds: Paleogeography and evolution of the
vertebrate fauna from the Balearic Islands
Pere Bover
a,
, Josep Quintana
b
, Josep Antoni Alcover
c
a
Division of Vertebrate Zoology/Mammalogy, American Museum of Natural History, Central Park West at 79th Street, New York, NY 10024, USA
b
Carrer Gustau Mas, 79, 1er, 07760 Ciutadella de Menorca, Illes Balears, Spain
c
Institut Mediterrani d’Estudis Avanc-ats, Ctra. Valldemossa km 7,5, 07122 Palma de Mallorca, Illes Balears, Spain
Available online 22 July 2007
Abstract
The Balearic Islands are an archipelago located in the Western Mediterranean Sea. Their isolation from the mainland allowed the
establishment of different faunas on each island. In Mallorca, the Pliocene fauna was composed of the so-called Myotragus-fauna
(mainly consisting of a bovid, a glirid and a soricid). In Menorca, it was constituted by the giant rabbit-fauna (mainly consisting of a
giant rabbit and a tortoise), and in the Pityusics by a tortoise, a lizard and two rodents. A main faunal turnover took place during the
Late Pliocene or Early Pleistocene: the Myotragus-fauna reached Menorca and replaced the giant rabbit fauna. In the Pityusics, all
mammals and the tortoise became extinct before the Late Pleistocene for unknown reasons, leaving birds and the lizard as the only
vertebrates of these islands. Almost all the endemic vertebrates of the Balearics became extinct probably due to the first human arrival to
the islands.
r2007 Elsevier Ltd and INQUA. All rights reserved.
1. Introduction
The Mediterranean is an almost completely closed sea
located between Europe and Africa, with a complex
geographical history. It contains archipelagos that differ
in area, geology, height above sea-level, isolation, anti-
quity, ecology and timing of human colonization (e.g.,
Maldonado, 1985;Margalef, 1985;Ramis and Alcover,
2004). Almost all the Mediterranean islands have very
important deposits of fossil vertebrates, some of which
have been known since the end of the 19th century. Thus,
the Mediterranean is an interesting scenario for the study
of island vertebrate evolution. The isolation of most of the
Mediterranean islands spreads over millions of years
(Myr), and fossil records have documented the presence
of very singular, highly modified species in these islands in
the past.
The main goal of this paper is to update the knowledge
of the vertebrate fossil fauna from the Balearic Islands
(sensu lato) since the start of their current isolation (i.e.,
since the last moment they were connected to the
surrounding mainland). The Balearic Islands are the most
isolated islands in the Mediterranean. This archipelago is
composed by two different groups of islands (‘‘sub-
archipelagos’’). The Western set, or Pityusic Islands,
consists of two main islands (Eivissa [‘‘Ibiza’’] and
Formentera) and nearly 60 smaller surrounding islets.
These two islands are separated by a narrow, shallow
channel, and they shared their faunas throughout their
paleogeographical history. They will be analyzed as a single
unit in this paper. On the other hand the Eastern set, or
Gymnesic Islands, is more isolated than the Pityusics and is
constituted by two main islands (Mallorca and Menorca)
and nearly 30 surrounding islets. These two islands display
a different paleogeographical history at the beginning of
their history as islands, and a peculiar faunal succession.
The current isolation of the Balearic Islands began at the
end of the Messinian salinity crisis (MSC), 5.3 Myr ago
(Gautier et al., 1994;Clauzon et al., 1996;Krijgsman et al.,
1999). Although some Middle and Late Miocene insular
faunas have been recorded, there is no evidence of their
continuity through the Messinian. The duration of the
ARTICLE IN PRESS
1040-6182/$ -see front matter r2007 Elsevier Ltd and INQUA. All rights reserved.
doi:10.1016/j.quaint.2007.06.039

Corresponding author. Tel.: +1 212 769 5693; fax: +1 212 769 5239.
E-mail addresses: pbover@amnh.org (P. Bover),
picoguevo@hotmail.com (J. Quintana),vieapba@uib.es (J.A. Alcover).

current isolation of these sub-archipelagos embraces
different climatic changes. On the one hand, the beginning
of the Late Pliocene coincides with the change from a
Subtropical climate [or warm-temperate (e.g., Fauquette
et al., 1999)] to a Mediterranean one (e.g., Shackleton
et al., 1984;Suc, 1984;Leroy and Dupont, 1994;Burkle,
1995;Bourillet et al., 2006). On the other hand, during the
last 2.5 million years the Mediterranean area has been
widely influenced by glaciations. The changing climate of
this area has influenced the paleogeography and evolution
of the species living there.
The Balearic Islands, due to their degree of isolation,
have been colonized by humans in a very recent time (in
comparison with less isolated Mediterranean islands).
Although until 2001 human arrival was assumed to have
occurred 8000 years ago (or even more) (e.g., Guerrero,
2000, 2001), some recent reviews document the first
evidence of human presence in Mallorca in the third
millennium BC (Alcover et al., 2001;Ramis et al., 2002),
and the human settlement occurred probably in the last
third of the millennium (Alcover, submitted).
Both the degree (distance from mainland) and duration
(millions of years) of isolation have been key factors for the
evolution of highly peculiar taxa in the Balearics. Thus, the
Balearic Islands have been considered ‘‘oceanic-like
islands’’ (Alcover et al., 1998). ‘‘Oceanic-like islands’’ were
described by these authors as islands that have been
previously connected to continents, but that are faunisti-
cally similar to oceanic islands (i.e., with a high level of
endemism and highly unbalanced, poor faunas), because
the island-continent connection occurred in a distant past,
was of short duration, or did not promote a complete
faunal transfer.
Although no direct evidence of the duration of the
Messinian connection of the Balearic Islands with the
surrounding mainlands is available, and there is no data
about how the geographic filtering took place, faunal
transfer has been very incomplete.
Both Balearic Islands groups contained highly distinctive
faunas in the Early Pliocene. They evolved to be the most
singular territories in the Mediterranean: the Pityusics were
the only islands in the whole Mediterranean Sea without
mammals during the Late Pleistocene and Early Holocene,
while the Gymnesics were the sole Mediterranean islands
inhabited by a highly modified caprine at that time
(Alcover et al., 1981).
2. Early insular faunas
Although the main goal of this paper is to focus on the
post-Messinian fossil faunas, a short sketch of the clearly
insular faunas known in the Balearic Islands predating the
Messinian crisis is presented here. The earliest faunas with
an unquestionable insular character known in the Balearic
Islands are represented by finds in four Miocene deposits
from Mallorca (Santa Margalida, Sant Llorenc-, Cova de
Cala Varques D, Cova des Coll) and two deposits from
Menorca (Punta Nati-2 and Es Cul de Sa Ferrada).
In Mallorca, the fauna from Santa Margalida-Sant
Llorenc-has been attributed to the Langhian age (MN 5;
Mein and Adrover, 1982), although a more recent attribu-
tion (Serravallian) cannot be rejected. The island character
of this fauna is demonstrated by three facts: it is
oligospecific, highly disharmonic, and endemic. The fauna
includes an ochotonid lagomorph (Gymnesicolagus gelaberti
Mein and Adrover 1982) and 3 glirid rodents (Carbomys
sacaresi Mein and Adrover 1982, Margaritamys llulli Mein
and Adrover 1982 and Peridyromys ordinasi Mein and
Adrover 1982) (Mein and Adrover, 1982;Adrover et al.,
1983, 1984). G. gelaberti is the largest of the known
Ochotonidae, and C. sacaresi is a giant glirid with
nonestablished taxonomic affinities. The recent exploration
of underwater caves excavated in the East Mallorcan
Miocene platform allowed the discovery of other Late
Miocene pre-Messinian deposits: Cova des Coll and Cova
de Cala Varques B (e.g., Gra
`cia et al., 2005). These deposits
have yielded remains of large terrestrial tortoises (Geoche-
lone sp), as well as remains of a sea turtle (Trionyx sp).
From Menorca, the deposit of Punta Nati 2 yielded
remains of a Gymnesicolagus, a glirid, similar to M. llulli
from the Mallorcan deposits of Santa Margalida and Sant
Llorenc-, and a large tortoise. Its precise chronology is
unknown, but it predates the Messinian. A small fragment
of a Gymnesicolagus jaw has been recovered from the
second Menorcan deposit, Es Cul de Sa Ferrada. This
deposit has been attributed to the Tortonian (Quintana
and Agustı
´, in press).
All these insular faunas suggest that in the Middle and
perhaps also in the Late Miocene there was a set of islands
near the coast of the Spanish mainland. Another insular
fauna from the Middle Miocene was found in the currently
mainland area of Murchas (Granada), Spain (Martin-
Sua
´rez et al., 1993). This fauna seems to be related to the
Gymnesicolagus faunas from Mallorca and Menorca, and it
contains some insular-evolved species descendants of
Pseudodryomys de Bruijn 1966 and Peridyromys Stehlin
and Schaub 1951.
3. Messinian salinity crisis
A key event affecting faunal distribution and evolution
in the Mediterranean during the Late Miocene was the
MSC. It promoted a significant faunal exchange between
Europe and Africa, and a faunal turnover in the Balearic
Islands (e.g., Agustı
´et al., 2006;van der Made et al., 2006
and references therein), and is considered to be the time of
colonization of the Balearic Islands by the fauna that
evolved in this territory during the Pliocene, and, in some
cases, until the Holocene (e.g., van der Made et al., 2006).
The MSC was discovered in the early 1970s (Hsu
¨et al.,
1973) and there is currently a general consensus on the age
that it took place, between 5.96 and 5.33 Myr (e.g.,
Krijgsman et al., 1999).
ARTICLE IN PRESS
P. Bover et al. / Quaternary International 182 (2008) 135–144136

The MSC occurred when the connection between the
Mediterranean Sea and the Atlantic Ocean was closed due
to tectonic displacements (Krijgsman et al., 1999), promot-
ing the partial (or perhaps total) Mediterranean dissecation
and a sea level drop of nearly 1500 m between 5.6 and
5.32 Myr (Clauzon et al., 1996). Currently, three different
faunal assemblages are recognized as living in the Balearics
since the MSC. Mallorca, Menorca and the Pityusics
display a remarkably different Pliocene vertebrate record
(Fig. 1).
4. Plio-Quaternary faunas
4.1. Mallorca
Mallorca is the sole Balearic Island with a clear
continuity in its terrestrial vertebrate fauna from the
Messinian to the Holocene. The land mammals present at
the time of human arrival are the direct descendants of
those that reached Mallorca during the Messinian (e.g.,
Bover and Alcover, 2003). The vertebrate fauna consists of
three land mammals, a lizard, two amphibians, bats and
birds.
The most remarkable taxon belonging to this fauna is
the bovid Myotragus Bate 1909. Its lineage evolved in a
highly peculiar way, and it is considered the key species to
understanding the palaeoecology of Mallorca, due to its
potential effect on the Mallorcan vegetation (e.g., Alcover
et al., 1999).
To date, five chronospecies have been described in this
bovid lineage. Alcover et al. (1981) claimed that the major
patterns recorded during its evolution consists of the
progressive reduction of the number of incisiform teeth
(incisors and canines) and premolars, the increase in the
degree of hypsondonty of all teeth, the acquisition of
evergrowing incisors, the shortening of the rostral part of
the skull and jaw, the lengthening of frontal bones, the
frontalization of eye sockets, the acquisition of short, stout
limb bones (with extremely short metapodials and pha-
langes), the progressive fusion of tarsal bones, the change
in the structure of the pelvis and the acquisition of a small
size. As a consequence of this insular evolution, each of the
Myotragus balearicus (Bate, 1909) bones is highly dis-
tinctive among caprines.
The acquisition process of these features can be traced
through the different species of the lineage, from
M. pepgonellae Moya
`-Sola
`and Pons-Moya
`1982 (attrib-
uted to the Early Pliocene), through M. antiquus (Pons-
Moya
`, 1977) (Middle Pliocene), M. kopperi Moya
`-Sola
`and
Pons-Moya
`1981 (Plio-Pleistocene boundary), Myotragus
batei Crusafont and Angel 1966 (Early-Middle Pleisto-
cene), to M. balearicus (uppermost part of the Middle
Pleistocene to the Holocene). The last three species are
characterized by the reduction of the number of incisors at
ARTICLE IN PRESS
Fig. 1. Diagram of the different faunal sets (only terrestrial vertebrates are figured) that lived in the Balearic Islands during the Plio-Quaternary.
(SF?) ¼Ses Fontanelles fauna, with unknown chronology (Late Miocene–Early Pliocene?). (*) ¼Considered as endemic fauna (M. binigausensis,
H. eliomyoides and N. meloussae, respectively) by Pons-Moya
`et al. (1981). The gray arrows show the contact between Menorca and Mallorca during
Quaternary glaciations. See text for further explanation.
P. Bover et al. / Quaternary International 182 (2008) 135–144 137

adult age, with a remarkably increased lengthening of their
enameled part.
M. balearicus presents a high number of anatomical
derived characters. Such apomorphies have been tradition-
ally interpreted as morphological adaptations to a Medi-
terranean xerophytic isolated environment free of
mammalian carnivores (e.g., Sondaar, 1977;Alcover et
al., 1981). Some of the peculiarities of M. balearicus are
related to an increase in the efficiency of herbivorous
feeding and, thus, to a higher potential impact on the
vegetation.
Caprines are considered some of the most efficient
consumers of vegetation, and the impact of goats on
islands is well known [see Campbell et al., 2004;Campbell
and Donlan, 2005, for a review of the known impacts of
goat introductions on islands]. Such a putative effect would
have been increased in the case of Mallorca and Menorca
where M. balearicus, the terminal species of the lineage
(Fig. 2), was not only the sole middle-sized herbivore, but
was also living in an isolated environment without
carnivore predators, and developed a highly powerful
feeding apparatus. Increased hypsodonty, reduction of
distal jaw length (Alcover et al., 1981), proportional
increase of insertion ridges for mastication muscles in skull
and jaw, and concentration effort mainly on M3 and distal
part of M2 (Bover, 2004) might be related to this high
efficiency of feeding on vegetation with a xerophytic
character.
The phylogenetic relationships of Myotragus are still not
clearly understood. Initially (Andrews, 1915;Gliozzi and
Malatesta, 1980;Palombo et al., 2006) it was related to
Nemorhaedus Smith 1827 and Capricornis Ogilby 1837.
These genera were traditionally related to Rupicapra de
Blainville 1816 and Oreamnos Rafinesque 1817 and
included with the former in Rupicaprini (see Simpson,
1945). Further approaches (e.g., Gentry, 1978;Gatesy et
al., 1997;Hassanin and Douzery, 1999, 2003) questioned
the monophily and recognition of the Rupicaprini and
Caprini by Simpson (1945). Recent biomolecular research
(Lalueza-Fox et al., 2005) supports a relationship between
Myotragus and Ovis Linnaeus 1857, excluding the assumed
relationship with the ‘‘Rupicaprini’’. A formerly proposed
relationship of Myotragus to the clade Ovis+Budorcas is
rejected as an artifact derived from a wrong adscription of
Budorcas Hodgson 1850 data in the GenBank (see Lalueza-
Fox et al., 2005).
Recently, Bover and Alcover (2005) proposed to include
the earlier Mallorcan Myotragus species in a new genus,
Insulotragus Bover and Alcover 2005. This proposal
emphasizes the existence of an evolutionary change
between the two earlier recognized species in the lineage
and the last three ones, although the new name could be
used as a subgenus of Myotragus to emphasize that they
belong to a single insular lineage. One of the most
distinctive characteristics of this bovid lineage is the
progressive reduction in the number of incisiform teeth.
This reduction has been interpreted as related to the
acquisition of monophiodoncy in these teeth (Bover and
Alcover, 1999).
The clearest sequence of the evolutionary patterns
explained above can be observed in the series extending
from M. kopperi (Plio-Pleistocene boundary), through
M. batei (Early-Middle Pleistocene) and M. balearicus
(latest Middle Pleistocene to Holocene). In the later
species, an extreme effect of insular evolution can be
observed, mainly on the feeding apparatus, locomotion,
sense organs and body size.
The fusion of some tarsal bones (naviculocuboid fused to
small and great cuneiforms and to the canon bone) was
first functionally described by Leinders and Sondaar
(1974), in which the zig-zag movements allowed by the
movement of these bones when unfused were not possible
in M. balearicus. Later, more features related to joint
stabilization and ‘‘low-gear’’ locomotion in the species
were described. Some of the most significant are the
presence of reduced joint angles in several bones (Spoor,
1988), the peculiar calcaneum shape (Moya
`-Sola
`, 1979),
the reduced metapodial joint surface (Spoor, 1988;Ko
¨hler,
1993), some metacarpal bone fusions (Bover et al., 2005)
and the femur, humerus–radius joint and pelvis character-
istics (Bover, 2004, 2006).
Ko
¨hler and Moya
`-Sola
`(2001) suggest that the presence
of notches in some M. balearicus phalange joints could be
related to the presence of intracapsular ligaments as a
special joint stabilization mechanism. Nevertheless, these
same notches are present on the phalange joints of some
recent bovids deposited in the National Museum of
Natural History, Smithsonian Institution, Washington,
DC. (NMNH). They correspond to very old specimens
from zoological parks. This suggests that the notches could
be more related to the acquisition of a very old age than to
ARTICLE IN PRESS
Fig. 2. M. balearicus skull (MNIB 81723) from Cova des Tancats
(Menorca) in lateral view. The presence of a single evergrowing incisor,
reduction of number of premolars and frontalization of eye orbits are
some of the most important derived features in the skull of the species.
Scale bar 2 cm.
P. Bover et al. / Quaternary International 182 (2008) 135–144138

the presence of a special joint stabilization mechanism
(Bover, 2004).
The proportional size of the brain is reduced in
M. balearicus (Ko
¨hler and Moya
`-Sola
`, 2004), and some
sense organs could have been less sensitive. It has been
suggested that the reduction of the eye sockets could have
affected the vision (Ko
¨hler and Moya
`-Sola
`, 2004) and the
presence of a less developed criba nasalis would have
affected the sense of smell (Bover and Tolosa, 2005).
Together with the ancestors of Myotragus, two other
terrestrial mammals evolved in Mallorca after the Messi-
nian, a glirid rodent, probably derived from Eliomys truci
Mein and Michaux 1970 or a close species, and a soricid
insectivore. Both genera evolved together with Myotragus
but they did not reach the extreme differentiation observed
in the bovid lineage. Two endemic genera were proposed
for them, Hypnomys Bate 1918 and Nesiotites Bate 1944,
sometimes included as subgenera of other genera (Eliomys
for the glirid and Episoriculus Horsfield 1851, Soriculus
Blyth 1854 or Asoriculus Kretzoi 1959 for the soricid).
These species are larger than their mainland relatives,
which is in agreement with the general patterns recorded
for island mammals (e.g., Thaler, 1973;Case, 1978;
Heaney, 1978).
Three species of Hypnomys (Rodentia: Gliridae) have
been described. Hypnomys waldreni Reumer 1979, which
coexisted with Myotragus antiquus (Middle Pliocene),
Hypnomys eliomyoides Agustı
´1980 [¼H. onicensis (Re-
umer 1994), sensu Reumer (1994)], contemporary with
Myotragus kopperi and probably M. batei (Plio-Quatern-
ary boundary to Middle Pleistocene) and Hypnomys
morpheus Bate 1918 which coexisted with M. balearicus
(uppermost part of the Middle Pleistocene to the Holocene)
(Alcover et al., 1981). Although a few glirid bones have
been found in the type deposit of Myotragus pepgonellae
(Cala Morlanda, Early Pliocene) (Bauza
´, 1961), they have
not been ascribed to any particular species (Alcover et al.,
1981).
The main evolutionary pattern observed in the Hyp-
nomys lineage consists of the progressive acquisition of a
large size, the increase of tooth crown height and the
acquisition of a flat occlusal surface in the molars (related
to the presence of xerophytic vegetation). According to
Mills (1976),H. morpheus was probably more adapted to a
rat life-style than to a dormouse life-style, probably
displaying a less scansorial behavior than recent dormice.
Only two species of Nesiotites (Soricomorpha: Soricidae)
have been described in Mallorca. Nesiotites hidalgoi Bate
1944 was described by Bate (1944) to include the fossil
shrew of Late Pleistocene deposits from Mallorca and
Menorca, coetaneous with M. balearicus and H. morpheus.
Reumer (1979) described its ancestor, N. ponsi Reumer
(1979), coming from deposits containing M. antiquus. This
species displays a small size and a characteristic dental
pattern, with secondary cusps on the M
1
and M
2
. Some
Soricidae remains described as cf. Nesiotites have been
obtained in the deposits where M. pepgonellae was present.
Furthermore, an intermediate form has been found in the
Pedrera de s’O
`nix deposit (from where M. kopperi was
recovered, Plio-Quaternary boundary) but it has been
described as Nesiotites aff. ponsi (Alcover et al., 1981).
Remains of a lizard have also been found in deposits
from the Early Pleistocene to the Holocene. Although the
Early Pleistocene remains were described as Podarcis sp,
and are characterized by a very small size, the Middle
Pleistocene to Holocene remains were ascribed to the same
species as is nowadays living in the isles surrounding
Mallorca and Menorca, Podarcis lilfordi (Gu
¨nther 1874).
Two amphibians are known from the Pleistocene
deposits of Mallorca. Discoglossus sp, a large-sized
discoglossid frog, is only known from Pedrera de s’O
`nix,
the type deposit of M. kopperi. It is absent in all the known
deposits of the Middle and Late Pleistocene. Alytes
muletensis Sanchı
´z and Adrover 1977, the Mallorcan
Midwife Toad, has been identified from deposits ranging
from the Early Pleistocene to the Holocene. After its
description as a fossil frog (Sanchı
´z and Adrover, 1977),
A. muletensis was found living in very inaccessible torrents
from the Serra de Tramuntana of Mallorca 25 years ago
(Mayol et al., 1981) and is now one of the main species
concerned in the wildlife preservation programs in the
Balearic Islands.
4.2. Menorca
Two successive post-Messinian insular faunas have been
reported from Menorca. The earliest faunal assemblage is
situated inside the Pliocene, and probably spread during
the Early and Middle Pliocene (Quintana, 1998;Alcover et
al., 1999). Its most characteristic elements are a giant
tortoise (Bate, 1914) and a yet unnamed giant rabbit
(Fig. 3). Additional taxa include the remains of a dormouse
(Muscardinus cyclopeus Agustı
´,Moya
`-Sola and Pons-
Moya
`1982), a bat (Rhinolophus cf. grivensis Depe
´ret
1892) (Pons-Moya
`et al., 1981) and several genera and
species of birds: a petrel (Pterodromoides minoricensis
Seguı
´, Quintana, Forno
´s and Alcover 2001), a woodcock
(Scolopax carmesinae Seguı
´1999), a crane (Camusia
quintanai Seguı
´2002), two owls [Tyto balearica Mourer-
Chauvire
´, Alcover, Moya
`-Sola
`and Pons-Moya
`1980 and
Athene sp (Seguı
´, 1998)], a crow (Corvus sp) and an
unidentified passeriform (Seguı
´, 1998).
Among the reptiles, fossils include a lizard (Podarcis sp)
(Bailo
´n, 2004), which could be the P. lilfordi ancestor), a
geckonid (Gekkonidae indet.), an amphisbaenid (Blanus
sp) (Garcia-Porta et al., 2002;Bailo
´n et al., 2005), a
colubrid (Coluber sp) (Bailo
´n et al., 2005), and two viperids
(Vipera natiensis Bailo
´n, Garcia-Porta and Quintana 2002
and Vipera sp) (Bailo
´n et al., 2002). Included in the giant
rabbit fauna there is just one species of amphibian: Latonia
sp (Quintana et al., 2005).
The low number of species, the lack of representatives of
Carnivora and Perissodactyla, the gigantism of the
dormouse and the rabbit, and the change in the morphol-
ARTICLE IN PRESS
P. Bover et al. / Quaternary International 182 (2008) 135–144 139

ogy of the skeleton of the latter, allow us to state that this
faunistic group evolved in island conditions. The giant
rabbit from Menorca clearly displays some of these insular
characteristics. It has been estimated to have had a body
weight of 14 kg, very small eye sockets and tympanic bulla,
a relatively small head in relation to the body, and limbs
and vertebral column shorter than those of the recent
cursorial rabbits (Quintana, 2005;Quintana et al., 2005).
A faunal turnover occurred in Menorca during the Late
Pliocene or Early Pleistocene. To date, the Menorca giant
rabbit fauna have not been recorded in Pleistocene
deposits, suggesting that this fauna disappeared from this
island before then. In Menorcan Pleistocene and Holocene
deposits only the Myotragus fauna is recorded. The arrival
of this fauna to Menorca could be related to the beginning
of the glaciations and the related melting of Mallorca and
Menorca (e.g., Cuerda, 1975).
Although a Myotragus species was described from the
Early Pleistocene of Menorca, Myotragus binigausensis
Moya
`-Sola
`and Pons-Moya
`1980, it has recently been
synonymized to M. batei (the same species as in Mallorca)
by Bover and Alcover (2000).M. binigausensis was initially
considered to be a representative of a different lineage of
the genus on the island of Menorca, whose ancestor
reached this island during the Messinian (Moya
`-Sola
`and
Pons-Moya
`, 1980;Alcover et al., 1981). Other endemic
mammalian species from Menorca that were described
from a Menorcan site, including the glirid H. eliomyoides
Agustı
´1980 and the soricid Nesiotites meloussae Pons-
Moya
`and Moya
`-Sola
`1980, are also currently considered
to be conspecific with the representative of those genera for
that period in Mallorca. Thus, the available evidence
argues in favor of the dispersal of these three land
mammals (Myotragus,Hypnomys and Nesiotites)to
Menorca from Mallorca during a glaciation period, at an
unknown moment in the Late Pliocene or Early Pleistocene
(Bover, 2004). No evidence of specific differentiation of any
mammal has been obtained in Menorca once the Myo-
tragus fauna reached the island, probably because of the
recurrent coalescence of both islands during the glacia-
tions, which allowed repetitive inbreeding between both
populations (Bover, 2004).
The lizard of genus Podarcis Wagler 1830 was also
present in Menorca, at least since the Pliocene (Bailo
´n,
2004). Its evolutionary lineage can be traced until the
arrival of the Romans (Reumer and Sanders, 1984).
Currently, as happens in Mallorca, the species survives
only on some small islets.
A species of Midwife toad, Alytes talaioticus Sanchı
´z and
Alcover 1982, was described in Menorca from Holocene
deposits (Sanchı
´z and Alcover, 1982). It is currently
considered to be the same species as the Mallorcan Alytes
muletensis (e.g., Barbadillo, 1987). A still undescribed
species of Discoglossus Otth 1837 is known from the Plio-
Pleistocene boundary deposit of Barranc de Binigaus, Sa
Segonya and Punta Esquitxador (Alcover et al., 1981;
Quintana, 1998).
4.3. Eivissa
An early insular fauna from Eivissa comes from Ses
Fontanelles. Its stratigraphic position is unknown. The Ses
Fontanelles fauna contains two bovids, two rodents (one
gerbilid, Protatera sp, and one glirid, Eliomys sp), an
insectivore, and a leporid (Alilepus sp) as well as some
reptiles (a lizard and a tortoise) (Moya
`-Sola
`et al., 1984,
1999). The presence of the two rodents and the leporid
suggest that this deposit could be attributed to the
Messinian (Agustı
´and Moya
`-Sola
`, 1990), given that these
taxa have also been recorded in deposits of the Late
Miocene and Early Pliocene from the Iberian Peninsula.
One of the bovids present in this deposit was identified as
Tyrrhenotragus sp by Agustı
´and Moya
`-Sola
`(1990).
Current knowledge, however, suggests that Tyrrhenotragus
Hu
¨rzeler and Engesser 1976 lived between 6 and 9 Myr ago,
and that the fauna of Ses Fontanelles has a more recent age
(Moya
`-Sola
`et al., 1999). Due to the paucity of more
material from the two bovids, it seems prudent to attribute
this fauna to the Late Miocene–Early Pliocene, namely in
the same compatible context as the giant rabbit fauna and
Myotragus fauna.
Only one Eivissian site is attributed here to the Late
Pliocene, Cova de Ca Na Reia. It is a karst deposit with a
ARTICLE IN PRESS
Fig. 3. Comparison between a right femur of the giant rabbit from
Menorca (a) and a right femur of a recent Oryctolagus cuniculus (b). The
difference in size, together with the different orientation and joint surface
of the femur head, are among the major distinctive features of the bone of
the Menorcan giant rabbit. Scale bar 2 cm.
P. Bover et al. / Quaternary International 182 (2008) 135–144140

fauna containing the remains of two glirids (Eivissia
canarreiensis Alcover and Agustı
´1985 and Hypnomys sp)
(Alcover and Agustı
´, 1985), a middle-sized tortoise
(Cheirogaster sp) (Bour, 1985) and a lizard (Podarcis sp)
(Kotsakis, 1981). Bats and birds are also present. This is
the type locality for Puffinus nestori Alcover 1989, the
presumed ancestor of Puffinus mauretanicus Lowe 1921/
yelkouan Acerbi 1827.
Giant tortoises are known from three other sites: Es
Poua
`s (lower levels), Pedrera de Can Bessora (Sant Antoni
de Portmany) and La Mola cliffs (Formentera). None of
these deposits has been isotopically dated and they should
be tentatively attributed to some time inside the Late
Pliocene–Middle Pleistocene. It is not known whether the
absence of glirid remains in these deposits represents only a
sampling problem or whether it reflects true absence.
Eight Late Pleistocene deposits are known from Eivissa
and Formentera. These deposits have yielded tens of
thousands of vertebrate bones (only one deposit, Es Poua
`s,
yielded over 120,000 bones). The vertebrate fauna consists
only of fossil birds, bats and a lizard species (Podarcis
pityusensis Bosca
´1883). The bird fauna is highly peculiar,
and has been ecologically paralleled to the Hawaiian fossil
bird fauna (Seguı
´and Alcover, 1999). One of the species of
this fauna is Rallus eivissensis McMinn, Palmer and
Alcover 2005, the sole endemic rail described so far from
a Mediterranean island (McMinn et al., 2005).
5. Faunal extinction
5.1. Last occurrence data
Key data for the establishment of the extinction
chronology of a species is the date of its last occurrence.
The last record of a species is an important datum because
it provides a terminus post quem for the extinction event. It
is also important to establish the earliest documentation of
the species absence (i.e., a datum representing a terminus
ante quem for the extinction event). The earliest age for the
absence of the endemic mammals was estimated through
the recurrent lack of evidence of their presence over all the
archaeological deposits containing bones dated previous to
2040 cal BC in Mallorca.
Currently, new chronological data directly obtained
from collagen samples of endemic mammal bones from
the Gymnesics is available (Bover and Alcover, 2003, and
Bover and Alcover, submitted). The available datings show
that Myotragus was present in Mallorca after 3700 cal BC
(although there is some discussion on this date: see van
Strydonck et al., 2005), later than 3970 cal BC in Menorca
(Quintana et al., 2003) and later than 3650 cal BC in
Cabrera (Bover and Alcover, 2003). Data available for
Mallorca document the presence of Hypnomys later than
4840 cal BC and Nesiotites later than 3030 cal BC (Bover
and Alcover, submitted). These are the datings of the last
occurrence records.
5.2. First human presence data
Currently, no direct evidence of the contact between the
first human settlers and the autochthonous fauna from the
Gymnesics has been recorded. The first archaeological
record showing unquestionable human presence in Mal-
lorca can be dated in the last century of the III millennium
BC (e.g., Alcover et al., 2001;Ramis et al., 2002). A more
recent analysis of the chronology suggests that the first
human arrival in Mallorca could have occurred at an
unknown moment between 2350 and 2150 BC, a date
highly compatible with the evidence available.
Although there is no direct evidence of contact between
humans and the autochthonous mammals from the
Gymnesics, the only explanation for the Myotragus
extinction appears to be related with human arrival. Even
in the absence of such evidence, the extinction of the
Quaternary mammal fauna is also probably related to
human presence.
Bover and Alcover (1999) situated the extinction of
Myotragus inside the range 3600–2030 BC. A new
approach on the accurate timing of the human colonization
(Alcover, submitted) suggests (but does not prove) that the
extinction occurred after 2350 cal BC. It represents the last
event of the extinction of an insular ‘‘megafauna’’ in the
Mediterranean. It probably occurred just after human
arrival, as well as occurred for the extinction of small
mammals (Bover and Alcover, submitted).
Extinction also affected the Pityusic endemic Rallus
eivissensis. The last evidence of its presence post-dates
5300 cal BC. Its extinction is presumably related to the
arrival of the first human settlers (McMinn et al., 2005).
The improvement of the palaeontological and archae-
ological record of the III millennium cal BC. in Mallorca
will be decisive to obtain more precise information of the
chronology and causes of the extinction of the endemic
fauna of the Balearic Islands.
6. Conclusion
The vertebrate fossil record in the Pliocene and
Quaternary of the Balearic Islands reveals that different
faunal successions took place on different islands, follow-
ing different paleogeographic events. In the Messinian all
the islands were connected in some way with the
surrounding mainland. Three different faunas inhabited
the islands at that time. The Myotragus fauna lived in
Mallorca, Menorca was inhabited by the giant rabbit fauna
and Eivissa and Formentera would have been inhabited by
a fauna that originated the Pliocene assembly of Cova de
Ca Na Reia, including two glirids and a giant tortoise.
Currently, it remains unknown whether three different
faunas colonized each island or whether the same faunal
assemblage reached all islands and became different after
local extinctions of several species. New studies of recently
discovered and known deposits would shed light on this
subject.
ARTICLE IN PRESS
P. Bover et al. / Quaternary International 182 (2008) 135–144 141

After the end of the MSC, these three faunas started to
evolve separately, in complete isolation from one another
and the mainland. Among the Messinian faunas only the
Mallorcan fauna survived in insular conditions until
human arrival. Although this fauna was initially exclusive
to Mallorca, it later spread to Menorca. It represents the
longest isolated evolutionary experiment in the Balearics
and one of its elements, Myotragus, represents an extreme
case of evolution under conditions of insularity.
Both in Menorca and Eivissa, some faunal succession
has been recorded. In Menorca, the faunal succession was
represented by a land mammal turnover. The giant rabbit
fauna was substituted by the Myotragus fauna. Although
the precise timing of this substitution is still not well
established, it should have occurred between the Middle
Pliocene and the Early Pleistocene, according to the faunal
record. The start of the glaciations 2.6–2.7 Myr ago (e.g.,
Shackleton et al., 1984, but see Mudelsee and Raymo,
2005), with the merging of Mallorca and Menorca, emerges
as a reasonable possibility. After the turnover, roughly the
same fauna lived in Mallorca and Menorca until human
arrival.
Faunal succession has also been recorded in Eivissa. A
part of the original fauna disappeared at an indeterminate
time between the Pliocene and the Late Pleistocene. This
faunal change took place without mammalian turnover.
Definitively, the three main Balearic Islands acted as
separate worlds after the MSC. Each island had its own
fauna, which evolved in isolation. The start of the
glaciations was probably the origin of the homogenization
of the Mallorcan and Menorcan faunas, through the
melting of the islands. In Eivissa, an unknown event was
the cause of a faunal change producing the complete
extinction of all terrestrial mammals. As a consequence of
all these changes, the Pityusics and the Gymnesics were
ecologically different at the time of the first human arrival.
Acknowledgments
The authors are indebted to Dr. Maria Rita Palombo
(Roma), for her invitation and patience. Dr. Adrian
Tejedor (New York) corrected the English of the manu-
script. One of the authors (PB) has a MEC—Fulbright
postdoctoral fellowship from the Secretarı
´a de Estado de
Universidades de Investigacio
´n of the Ministerio de
Educacio
´n y Ciencia of Spain. This paper is included in
the Research Project CGT 2004-04612 of the Spanish
MEC.
References
Adrover, R., Agustı
´, J., Moya
`-Sola
`, S., Pons-Moya
`, J., 1983–1984. Nueva
localidad de micromamı
´feros insulares del Mioceno medio en las
proximidades de San Lorenzo en la isla de Mallorca. Paleontologia i
Evolucio
´18, 121–129.
Agustı
´, J., 1980. Hypnomys eliomyoides nov. sp., nuevo glı
´rido (Rodentia,
Mammalia) del Pleistoceno de Menorca (Islas Baleares). Endins 7,
49–52.
Agustı
´, J., Moya
`-Sola
`, S., 1990. Neogene—Quaternary mammalian
faunas of the Balearics. In: Azzaroli, A. (Ed.), Biogeographical
Aspects of Insularity, vol. 238. Atti dei Convegni Lincei, Accademia
Nazionale dei Lincei, Roma, Italy, pp. 459–469.
Agustı
´, J., Moya
`-Sola
`, S., Pons-Moya
`, J., 1982. Une espe
`ce ge
´ante de
Muscardinus Kaup, 1829 (Gliridae, Rodentia, Mammalia) dans le
gisement karstique de Cala es Pou (Mioce
`ne supe
´rieur de Minorque,
Bale
´ares). Geobios 15, 783–789.
Agustı
´, J., Garce
´s, M., Krijgsman, W., 2006. Evidence for African–Iberian
exchanges during the Messinian in the Spanish mammalian record.
Palaeogeography, Palaeoclimatology, Palaeoecology 238, 5–14.
Alcover, J.A., 1989. Les aus fo
`ssils de la Cova de ca Na Reia. Endins
14–15, 95–100.
Alcover, J.A., submitted. Disentangling the mallorcan first known settlers’
cultural identity, contacts, source region and precise colonization
chronology. Journal of World Prehistory.
Alcover, J.A., Agustı
´, J., 1985. Eliomys (Eivissia) canarreiensis n. sgen. n.
sp.; nou Glirid del Pleistoce de la cova de Ca Na Reia (Pitiuses).
Endins 10–11, 51–56.
Alcover, J.A., Moya
`-Sola
`, S., Pons-Moya
`, J., 1981. Les quimeres del
passat. In: Els vertebrats fo
`ssils del Plio-Quaternari de les Balears i
Pitiu
¨ses, Monografies Cientı
´fiques, 1, Editorial Moll, Palma de
Mallorca, pp. 1–260.
Alcover, J.A., Sans, A., Palmer, M., 1998. The extent of extinctions of
mammals on islands. Journal of Biogeography 25, 913–918.
Alcover, J.A., Bover, P., Seguı
´, B., 1999. Paleoecologia de les illes. In:
Alcover, J.A. (Ed.), Ecologia de les Illes, Monografies de la Societat
d’Histo
`ria Natural de les Balears, 6/Monografia de l’institut d’Estudis
Balea
`rics, 66, Palma de Mallorca, Spain, pp. 169–204.
Alcover, J.A., Ramis, D., Coll, J., Trias, M., 2001. Bases per al
coneixement del contacte entre els primers colonitzadors humans i la
naturalesa de les Balears. Endins 24, 5–57.
Andrews, C.W., 1915. A description of the skull and skeleton of a
peculiarly modified rupicaprine antelope (Myotragus balearicus Bate
1909), with a notice of a new variety, M. balearicus var. major.
Philosophical Transactions of the Royal Society, London ser. B 206,
281–305.
Bailo
´n, S., 2004. Fossil records of lacertidae in Mediterranean islands: the
state of the art. In: Pe
´rez-Mellado, V., Riera, N., Perera, A. (Eds.),
Proceedings of the Fourth International Symposium of the Lacertids
of the Mediterranean Basin, 12, Institut Menorquı
´d’Estudis, Mao
´, pp.
37–62.
Bailo
´n, S., Garcı
´a-Porta, J., Quintana-Cardona, J., 2002. Premie
`re
de
´couverte de Viperidae (Reptilia, Serpentes) dans les ıˆles Bale
´ares
(Espagne): des vipe
`res du Ne
´oge
`ne de Minorque. Description d’une
nouvelle espe
`ce du Plioce
`ne. Comptes Rendus Palevol 1, 227–234.
Bailo
´n, S., Quintana, J., Garcia-Porta, J., 2005. Primer registro fo
´sil de las
familias Gekkonidae (Lacertilia) y Colubridae (Serpentes) en el
Plioceno de Punta Nati (Menorca, Islas Baleares). In: Alcover, J.A.,
Bover, P. (Eds.), Proceedings of the International Symposium Insular
Vertebrate Evolution: The Palaeontological Approach, Monografies
de la Societat d’Histo
`ria Natural de les Balears, vol. 12. Palma de
Mallorca, Spain, pp. 27–32.
Barbadillo, L.J., 1987. La Guı
´a de Incafo de los anfibios y reptiles de la
Penı
´nsula Ibe
´rica, Islas Baleares y Canarias. Incafo, Madrid.
Bate, D.M.A., 1909. Preliminary note on a new artiodactyle from
Mallorca Myotragus balearicus, gen. et sp. nov. Geological Magazine
December 5 (6), 385–389.
Bate, D.M.A., 1914. A gigantic land tortoise from the Pleistocene of
Menorca. Geological Magazine December 6 (1), 100–107.
Bate, D.M.A., 1918. On a new genus of extinct muscardine rodent from
the Balearic islands. In: Proceedings of the Zoological Society London,
pp. 209–222.
Bate, D.M.A., 1944. Pleistocene shrews from the larger Western
Mediterranean islands. Annals and Magazine of Natural History
series 11–16, 738–769.
Bauza
´, J., 1961. Contribuciones a la geologı
´a de Mallorca. Boletı
´ndela
Sociedad de Historia Natural de las Baleares 7, 31–35.
ARTICLE IN PRESS
P. Bover et al. / Quaternary International 182 (2008) 135–144142

Bosca
´, E., 1883. Exploracion herpetolo
´gica de la Isla de I
´biza. Anales de la
Sociedad Espan
˜ola de Historia Natural 12, 241–250.
Bour, R., 1985. Una nova tortuga terrestre del Pleistoce
´d’Eivissa: la
tortuga de la cova de Ca Na Reia. Endins 10–11, 57–62.
Bourillet, J.F., Zaragosi, S., Mulder, T., 2006. The French Atlantic margin
and deep-sea submarine systems. Geo-Marine Letters 26, 311–315.
Bover, P., 2004. Noves aportacions al coneixement del ge
`nere Myotragus
Bate, 1909 (Artiodactyla, Caprinae) de les Illes Balears. Ph.D Thesis.
Universitat de les Illes Balears, Palma de Mallorca.
Bover, P., 2006. New morphological data for Myotragus balearicus
(Artiodactyla, Caprinae), from the Balearic Islands (Western Medi-
terranean. Journal of Vertebrate Paleontology 26 Abstracts, 44A.
Bover, P., Alcover, J.A., 1999. The evolution and ontogeny of the
dentition of Myotragus balearicus Bate 1909 (Artiodactyla: Caprinae):
evidences from new fossil data. Biological Journal of the Linnean
Society 68, 401–428.
Bover, P., Alcover, J.A., 2000. La identitat taxono
`mica de Myotragus
binigausensis Moya
`-Sola
`i Pons-Moya
`1980. Endins 23, 83–88.
Bover, P., Alcover, J.A., 2003. Understanding late Quaternary extinctions:
the case of Myotragus balearicus Bate 1909. Journal of Biogeography
30, 711–721.
Bover, P., Alcover, J.A., 2005. A taxonomic approach to the insular
caprines of the Gymnesic Islands (western Mediterranean Sea).
Quaternaire, Hors-Se
´rie 2, 213–220.
Bover, P., Alcover, J.A., submitted. Extinction of the autochthonous small
mammals from Mallorca (Gymnesic Islands, Western Mediterranean
Sea) and its ecological consequences. Journal of Biogeography.
Bover, P., Tolosa, F., 2005. The olfactory ability of Myotragus balearicus:
preliminary notes. In: Alcover, J.A., Bover, P. (Eds.), Proceedings of
the International Symposium Insular Vertebrate Evolution: the
Palaeontological Approach, vol. 12, Monografies de la Societat
d’Histo
`ria Natural de les Balears. Palma de Mallorca, Spain, pp.
85–94.
Bover, P., Forno
´s, J.J., Alcover, J.A., 2005. Carpal bones, carpal fusions
and footprints of Myotragus: clues for locomotion and behavior. In:
Alcover, J.A., Bover, P. (Eds.), Proceedings of the International
Symposium Insular Vertebrate Evolution: the Palaeontological Ap-
proach, Monografies de la Societat d’Histo
`ria Natural de les Balears,
vol. 12., Palma de Mallorca, Spain, pp. 59-72
Burkle, L.H., 1995. Current issues in Pliocene paleoclimatology.
In: Vrba, E.S., Denton, G.H., Partridge, T.C., Burkle, L.H. (Eds.),
Paleoclimate and Evolution with Emphasis on Human Origins. Yale
University Press, New Haven, CT, pp. 3–7.
Campbell, K., Donlan, C.J., 2005. Feral goat eradications on islands.
Conservation Biology 19, 1362–1374.
Campbell, K., Donlan, C.J., Cruz, F., Carrion, V., 2004. Eradication of
feral goats Capra hircus from Pinta Island, Gala
´pagos, Ecuador. Oryx
38, 328–333.
Case, T.J., 1978. A general explanation for insular body size trends in
terrestrial vertebrates. Ecology 59, 1–18.
Clauzon, G., Suc, J-P., Gautier, F., Berger, A., Loutre, M-F., 1996.
Alternate interpretation of the Messinian salinity crisis: controversy
resolved? Geology 24, 363–366.
Crusafont, M., Angel, B., 1966. Un Myotragus (Mammife
`re Ruminant),
dans le Villafranchien de l’ıˆle de Majorque: Myotragus batei, nov. sp.
Comptes Rendues de l’Academie de Sciences Paris 262, 2012–2014.
Cuerda, J., 1975. Los tiempos cuaternarios en Baleares. Edit. Institut
Estudis Balea
`rics, Palma de Mallorca.
Fauquette, S., Suc, J.P., Guiot, J., Diniz, F., Feddi, N., Zheng, Z.,
Bessais, E., Drivaliari, A., 1999. Climate and biomes in the West
Mediterranean area during the Pliocene. Palaeogeography, Palaeocli-
matology, Palaeoecology 152, 15–36.
Garcia-Porta, J., Quintana, J., Bailo
´n, S., 2002. Primer hallazgo de Blanus
sp. (Amphisbaenidae, Reptilia) en el Neo
´geno balear. Revista
Espan
˜ola de Herpetologı
´a 16, 19–28.
Gatesy, J., Amato, G., Vbra, E., Schaller, G., Desalle, R., 1997. A cladistic
analysis of mithocondrial ribosomal DNA from the Bovidae.
Molecular Phylogenetics and Evolution 7, 303–319.
Gautier, F., Clauzon, G., Suc, J.P., Cravatte, J., Violanti, D., 1994. Age et
dure
´e de la crise de salinite
´Me
´ssinienne. Comptes Rendues de
l’Academie de Sciences Paris, ser. II 318, 1103–1109.
Gentry, A.W., 1978. Bovidae. In: Maglio, V.J., Cooke, H.B.S. (Eds.),
Evolution of African Mammals. Harvard University Press, Cam-
bridge, London, pp. 540–572.
Gliozzi, E., Malatesta, A., 1980. The Quaternary goat of Capo Figari
(Northeastern Sardinia). Geologica Romana 19, 295–347.
Gra
`cia, F., Clamor, B., Jaume, D., Forno
´s, J.J., Uriz, M.J., Martin, D.,
Gil, J., Gracia, P., Febrer, M., Pons, G., 2005. La cova des Coll
(Felanitx, Mallorca): espe
`leoge
`nesi, geomorfologia, hidrologia, sedi-
mentologia, fauna i conservacio
´. Endins 27, 141–186.
Guerrero, V.M., 2000. La colonizacio
´n humana de Mallorca en el
contexto de las islas occidentales del Mediterra
´neo: una revisio
´n
crı
´tica. In: Guerrero, V.M., Gorne
´s, S. (Coord.), Colonizacio
´n humana
en ambientes insulares. Universitat de les Illes Balears, Palma de
Mallorca, pp. 99–190.
Guerrero, V.M., 2001. The Balearic islands: prehistoric colonization of the
furthest Mediterranean islands from the mainland. Journal of
Mediterranean Archaeology 14, 136–157.
Gu
¨nther, A., 1874. Description of a new European species of Zootoca.
Annals and Magazine of Natural History, August.
Hassanin, A., Douzery, E.J.P., 1999. The tribal radiation of the family
Bovidae (Artiodactyla) and the evolution of the mitochondrial
Cytochrome b gene. Molecular Phylogenetics and Evolution 13,
227–243.
Hassanin, A., Douzery, E.J.P., 2003. Molecular and morphological
phylogenies of Ruminantia and the alternative position of the
Moschidae. Systematic Biology 52, 206–228.
Heaney, L.R., 1978. Island area and body size of insular mammals:
evidence from tri-colored squirrel (Callosciurus prevosti) of Southeast
Asia. Evolution 32, 29–44.
Hsu
¨, K.J., Cita, M.B., Ryan, W.B.F., 1973. Late Miocene dessication of
the Mediterranean. Nature 242, 240–244.
Ko
¨hler, M., 1993. Skeleton and habitat of recent and fossil ruminants.
Mu
¨nchen Geowissenchaftliche Abhanlundgen A 25, 1–88.
Ko
¨hler, M., Moya
`-Sola
`, S., 2001. Phalangeal adaptations in the fossil
insular goat Myotragus. Journal of Vertebrate Paleontology 21,
621–624.
Ko
¨hler, M., Moya
`-Sola
`, S., 2004. Reduction of brain size reduction and
sense organs in the fossil insular bovid Myotragus. Brain, Behavior and
Evolution 63, 125–140.
Kotsakis, T., 1981. Le lucertole (Lacertidae, Squamata) del Pliocene,
Pleistocene e Olocene delle baleari. Bolletı
´de la Societat d’Histo
`ria
Natural de les Balears 25, 135–150.
Krijgsman, W., Hilgen, F.J., Raffi, I., Sierro, F.J., Wilson, D.S., 1999.
Chronology, causes and progression of the Messinian salinity crisis.
Nature 400, 652–655.
Lalueza-Fox, C., Castresana, C., Sampietro, L., Marque
`s-Bonet, T.,
Alcover, J.A., Bertranpetit, J., 2005. Molecular dating of caprines
using ancient DNA sequences of Myotragus balearicus, an extinct
endemic Balearic mammal. BMC Evolutionary Biology 5, 70.
Leinders, J.J.M., Sondaar, P.Y., 1974. On functional fusions in footbones
of ungulates. Zeitschrift fu
¨rSa
¨ugetierkunde 39, 109–115.
Leroy, S., Dupont, L., 1994. Development of vegetation and continental
aridity in northwestern Africa during the late Pliocene: the pollen
record of ODP Site 658. Palaeogeography, Palaeoclimatology,
Palaeoecology 109, 295–316.
Maldonado, A., 1985. Evolution of the Mediterranean basins and a
reconstruction of the Cenozoic palaeoceanography. In: Margalef, R.
(Ed.), Western Mediterranean. Pergamon Press, London, pp. 18–61.
Margalef, R. (Ed.), 1985. Western Mediterranean. Pergamon Press,
London.
Martin-Sua
´rez, M., Freudenthal, M., Agustı
´, J., 1993. Micromammals of
the middle Miocene of the Granada basin (Spain). Geobios Me
´moire
Spe
´cial 6, 451–463.
Mayol, J., Alcover, J.A., Alomar, G., Pomar, G., Jurado, J., Jaume, D.,
1981. Supervive
`ncia de Baleaphryne (Amphibia: Anura: Discoglossi-
ARTICLE IN PRESS
P. Bover et al. / Quaternary International 182 (2008) 135–144 143

dae) a les muntanyes de Mallorca. Nota preliminar. Butlletı
´de la
Institucio
´Catalana d’Histo
`ria Natural 45 (secc. Zool. 3), pp. 115–119.
McMinn, M., Palmer, M., Alcover, J.A., 2005. A new species of rail (Aves:
Rallidae) from the Late Pleistocene and Holocene of Eivissa (Pityusic
Islands, western Mediterranean). Ibis 147, 706–716.
Mein, P., Adrover, R., 1982. Une faunule de mammife
`res insulares dans le
Mioce
`ne moyen de Majorque (Iles Bale
´ares). Geobios 6, 405–463.
Mills, D.H., 1976. Osteological study of the Pleistocene dormouse
Hypnomys morpheus Bate from Mallorca (Rodentia, Gliridae).
Publications of the Paleontological Institution of the University of
Uppsala, 4, pp. 5–73.
Mourer-Chauvire
´, C., Alcover, J.A., Moya
`-Sola
`, S., Pons-Moya
`, J., 1980.
Une nouvelle forme insulaire d’effraie geante, Tyto balearica n. sp.,
(Aves, Strigiformes), du Plio-Pleistocene des Baleares. Geobios 13,
803–811.
Moya
`-Sola
`, S., 1979. Morfologı
´a funcional del tarso en el ge
´nero
Myotragus Bate, 1909 (Artiodactyla, Rupicaprini). Acta Geologica
Hispanica 3, 87–91.
Moya
`-Sola
`, S., Pons-Moya
`, J., 1980. Una nueva especie del ge
´nero
Myotragus Bate, 1909 (Mammalia, Bovidae) en la isla de Menorca:
Myotragus binigausensis nov.sp. Implicaciones paleozoogeogra
´ficas.
Endins 7, 37–47.
Moya
`-Sola
`, S., Pons-Moya
`, J., 1981. Myotragus kopperi, une nouvelle
espe
`ce de Myotragus Bate 1909 (Mammalia, Artiodactyla, Rupicapri-
ni). In: Proceedings of the Koninklijke Nederlandse Akademie van
Wetenschappen ser. B, 84, pp. 57-69.
Moya
`-Sola
`, S., Pons-Moya
`, J., 1982. Myotragus pepgonellae nov. sp. un
primitivo representante del ge
´nero Myotragus Bate, 1909 (Bovidae,
Mammalia) en la isla de Mallorca (Baleares). Acta Geologica
Hispanica 12, 77–87.
Moya
`-Sola
`, S., Pons-Moya
`, J., Alcover, J.A., Agustı
´, J., 1984. La fauna de
vertebrados neo
´geno-cuaternaria de Eivissa (Pitiuses). Nota prelimi-
nar. Acta Geologica Hispanica 19, 33–35.
Moya
`-Sola
`, S., Quintana, J., Alcover, J.A., Ko
¨hler, M., 1999. Endemic
island faunas of the Mediterranean Miocene. In: Ro
¨ssner, G.E.,
Heissig, K. (Eds.), The Miocene Land Mammals of Europe. Pfeil,
Munchen, pp. 435–442.
Mudelsee, M., Raymo, M.E., 2005. Slow dynamics of the Northern
Hemisphere glaciation. Paleoceanography 20, PA4022.
Palombo, M.R., Bover, P., Valli, A.M.F., Alcover, J.A., 2006. The Plio-
Pleistocene endemic bovids from the Western Mediterranean islands:
knowledge, problems and perspectives. Hellenic Journal of Geos-
ciences 41, 153–162.
Pons-Moya
`, J., 1977. La nouvelle espe
`ce Myotragus antiquus de l’ıˆ le de
Majorque (Bale
´ares). In: Proceedings of the Koninklijke Nederlandse
Akademie van Wetenschappen Ser. B 80, pp. 215–221.
Pons-Moya
`, J., Moya
`-Sola
`, S., 1980. Nuevo representante del ge
´nero
Nesiotites Bate 1944; Nesiotites meloussae nov.sp. (Insectivora,
Soricidae) de los rellenos ca
´rsticos del Barranc de Binigaus (Es
Mercadal, Menorca). Endins 7, 53–56.
Pons-Moya
`, J., Moya
`-Sola
`, S., Agustı
´, J., Alcover, J.A., 1981. La fauna de
mamı
´feros de los yacimientos menorquines con Geochelone gymnesica
(Bate, 1914). Nota Preliminar. Acta Geologica Hispanica 16, 129–130.
Quintana, J., 1998. Aproximacio
´n a los yacimientos de vertebrados del
Mio-Pleistoceno de la isla de Menorca. Bolletı
´de la Societat d’Histo
`ria
Natural de les Balears 41, 101–117.
Quintana, J., 2005. Estudio morfolo
´gico y funcional de Nuralagus rex.
Ph.D. Thesis. Universitat Auto
`noma de Barcelona, Barcelona.
Quintana, J., Agustı
´, J., In press. Los mamı
´feros del Mioceno medio y
superior de Menorca (Islas Baleares, Mediterra
´neo occiental). Geo-
bios.
Quintana, J., Bover, P., Ramis, D., Alcover, J.A., 2003. Cronologia de la
desaparicio
´de Myotragus balearicus Bate 1909 a Menorca. Endins 25,
155–158.
Quintana, J., Moya
`, S., Ko
¨hler, M., 2005. El conejo gigante de los
depo
´sitos ca
´rsticos de Punta Nati—Cala’s Pous (Menorca, Illes
Balears). In: Alcover, J.A., Bover, P. (Eds.), Proceedings of the
International Symposium Insular Vertebrate Evolution: the Palaeon-
tological Approach, Monografies de la Societat d’Histo
`ria Natural de
les Balears, Palma de Mallorca, Spain, 12, pp. 297–308.
Ramis, D., Alcover, J.A., 2004. Irrupcio
´n humana y extincio
´n faunı
´stica
en las grandes islas del Mediterra
´neo durante el Holoceno. In:
Baquedano, E., Rubio Jara, S. (Eds.), Miscela
´nea en Homenaje a
Emiliano Aguirre, vol. IV. Arqueologı
´a. Museo Regional de la
Comunidad de Madrid, Alcala
´de Henares, pp. 390–401.
Ramis, D., Alcover, J.A., Coll, J., Trias, M., 2002. The chronology of the
first settlement of the Balearic Islands. Journal of Mediterranean
Archaeology 15, 3–24.
Reumer, J.W.F., 1979. On two new micromammals from the Pleistocene
of Mallorca. In: Proceedings of the Koninklijke Nederlandse
Akademie van Wetenschappen, ser. B 82, pp. 473–482.
Reumer, J.W.F., 1994. Eliomys (Hypnomys) onicensis nomen novum, to
replace the homonym Hypnomys intermedius Reumer, 1981 (Rodentia:
Gliridae) from Majorca. Zeitschrift fu
¨rSa
¨ugetierkunde 59, 380–381.
Reumer, J.W.F., Sanders, E.A., 1984. Changes in the vertebrate fauna of
Menorca in prehistoric and classical times. Zeitschrift fu
¨rSa
¨ugetier-
kunde 49, 321–325.
Sanchı
´z, F.B., Adrover, R., 1977. Anfibios fo
´siles del Pleistoceno de
Mallorca. Don
˜ana Acta Vertebrata 4, 5–25.
Sanchı
´z, F.B., Alcover, J.A., 1982. Un nou discoglo
`ssid (Amphibia:
Anura) de l’Holoce
`de Menorca. Butlletı
´de la Institucio
´Catalana
d’Histo
`ria Natural 48, 99–105.
Seguı
´, B., 1998. Els ocells fo
`ssils de Mallorca i Menorca. Successio
´
estratigra
`fica d’aus en els rebliments ca
`rstics de les Gimne
`sies. Ph.D.
Thesis. Universitat de les Illes Balears, Palma de Mallorca.
Seguı
´, B., 1999. A late tertiary woodcock from Menorca, Balearic islands,
Western Mediterranean. The Condor 101, 909–915.
Seguı
´, B., 2002. A new genus of crane (Aves: Gruiformes) from the late
tertiary of the Balearic islands, Western Mediterranean. Ibis 144,
411–422.
Seguı
´, B., Alcover, J.A., 1999. Comparison of ecological patterns in
insular bird faunas: a case study from the western Mediterranean and
Hawaii. Smithsonian Contributions to Palaeobiology 89, 67–73.
Seguı
´, B., Quintana, J., Forno
´s, J., Alcover, J.A., 2001. A new genus of
fulmarine petrel (aves: Procellariiformes) from the Late Miocene of
Menorca, Western Mediterranean. Paleontology 44, 933–948.
Shackleton, N.J., Backman, J., Zimmerman, H., Kent, D.V., Hall, M.A.,
Roberts, D.G., Schnitker, D., Baldauf, J.G., Desprairies, A.,
Homrighausen, R., Huddlestun, P., Keene, J.B., Kaltenback, A.J.,
Krumsiek, K.A.O., Morton, A.C., Murray, J.W., Westberg-Smith, J.,
1984. Oxygen isotope calibration of the onset of ice-rafting and history
of glaciation in the North Atlantic region. Nature 307, 620–623.
Simpson, G.G., 1945. The principles of the classification and a
classification of mammals. Bulletin of the American Museum of
Natural History 85, 1–350.
Sondaar, P.Y., 1977. Insularity and its effects on mammal evolution. In:
Hecht, M.K., Goody, P.C., Hecht, B.M. (Eds.), Major Patterns in
Vertebrate Evolution. Plenum Publishing Corporation, New York, pp.
671–707.
Spoor, C.F., 1988. The limb bones of Myotragus balearicus Bate, 1909. In:
Proceedings of the Koninklijke Nederlandse Akademie van We-
tenschappen, ser. B 91, pp. 295–308.
Suc, J.P., 1984. Origin and evolution of the Mediterranean vegetation and
climate in Europe. Nature 307, 429–432.
Thaler, L., 1973. Nanisme et gigantisme insulaires. La Recherche 37,
741–750.
van der Made, J., Morales, J., Montoya, P., 2006. Late Miocene turnover
in the Spanish mammal record in relation to palaeoclimate and the
Messinian salinity crisis. Palaeogeography, Palaeoclimatology, Pa-
laeoecology 238, 228–246.
van Strydonck, M., Boudin, M., Ervynck, A., 2005. Humans and
Myotragus: the issue of sample integrity in radiocarbon dating. In:
Alcover, J.A., Bover, P. (Eds.), Proceedings of the International
Symposium Insular Vertebrate Evolution: the Palaeontological Ap-
proach, Monografies de la Societat d’Histo
`ria Natural de les Balears,
Palma de Mallorca, Spain, 12, pp. 369–376.
ARTICLE IN PRESS
P. Bover et al. / Quaternary International 182 (2008) 135–144144