Fossil seeds from the pliocene of Menorca and Eivissa : Balearic Islands, Western Mediterranean
ENDINS, núm. 27. 2005. Mallorca
FOSSIL SEEDS FROM THE PLIOCENE OF MENORCA AND
EIVISSA (BALEARIC ISLANDS, WESTERN MEDITERRANEAN)
by Anna TRAVESET 1, Josep QUINTANA 2 and Josep Antoni ALCOVER 3
Abstract
The presence of Cneorum tricoccon and Crataegus monogyna during the Plio-
cene of Menorca, and of Cneorum tricoccon and Pistacia lentiscus during the Plio-
cene of Eivissa is documented from fossil seeds obtained in karstic deposits. We
compare the size of abundant fossil Cneorum seeds from Menorca with those obtai-
ned from the only extant population in this island. The finding of such Cneorum
seeds demonstrates the wider distribution of this species in the past in Menorca,
before the disappearance of its main disperser, the endemic lizard Podarcis lilfordi
(Lacertidae).
Resum
Es documenta la presència de Cneorum tricoccon i de Crataegus monogyna al
Pliocè de Menorca, i de Cneorum tricoccon i de Pistacia lentiscus al Pliocè d’Eivis-
sa a partir de les llavors fòssils obtingudes a dipòsits càrstics. Es compara la mida
dels fòssils de Cneorum tricoccon amb la de les llavors de la població actual de
l’illa de Menorca. La presència d’aquestes llavors fòssils de Cneorum a Menorca
demostra la seva major distribució en el passat, abans de la desaparació del seu
principal dispersor, la sargantana balear Podarcis lilfordi (Lacertidae).
Introduction
The vegetal macrofossil record of the Balearic
BAUZÀ (1971) whereas another study dealt with the vas-
Islands has received little attention compared to the ani-
cular flora of the Primary (BOURROUILH, 1973; LLOM-
mal component, mainly mammals, reptiles and birds
PART et al., 1979). From Menorca, the only studies that
(e.g., ALCOVER et al., 1981; BOUR, 1985; AGUSTÍ and
mention the presence of vegetal macrofossils (Cneorum
MOYÀ-SOLÀ, 1990; ALCOVER and McMINN, 1992;
tricoccon seeds) are QUINTANA (1998a,b) and QUINTA-
SONDAAR et al., 1995; SEGUÍ , 1999; BOVER, 2004;
NA et al. (2004) who found them in a couple of deposits
KÖHLER & MOYÀ-SOLÀ, 2004; QUINTANA et al., 2004;
of the north-eastern coast (deposits 11 and 28 of QUIN-
QUINTANA, 2005). Until the last decade, most informa-
TANA, 1998a). More recent information about the vege-
tion on palaeobotany of these islands came from old stu-
tational history of Mallorca and Menorca derives from
dies, mostly performed in Mallorca, on material coming
palynological studies performed during the last decade
from Oligocene and early Miocene (Burdigalian) depo-
(e.g., YLL et al., 1999, and references therein).
sits, which allowed the description of several new plant
In this paper, we present data on fossil seeds that
species (e.g., DEPAPE and FALLOT, 1928; ARENES,
have been obtained in two karstic deposits attributed to
1951; ARENES and DEPAPE, 1954; BAUZÀ, 1956;
Pliocene, one in Menorca and another in Eivissa, repre-
COLOM, 1982; RAMOS & ÁLVAREZ, 1989-90; HABLY &
senting the first fossil record of Neogene seeds in the
FERNÁNDEZ MARRON, 1998). The Triassic flora of
Balearic Islands. In the first case, the finding of a large
Mallorca was also examined several decades ago by
number of seeds from the same species (C. tricoccon)
allowed to compare their dimensions with those from
current seeds collected at the only extant population in
1
Institut Mediterrani d’Estudis Avançats (CSIC-UIB), Cr. Miquel
the island, in Cala Mesquida. Such fossil seeds eviden-
Marqués 21, 07190 Esporles, Mallorca, Illes Balears, Spain
2
Institut de Paleontologia Miquel Crusafont, C/ Escola Industrial 23,
ce that this shrub has become extinct from areas where
08201 Sabadell, Barcelona, Catalonia, Spain; current address: C/
it used to live in the past. In the case of Eivissa, fossil
Gustavo Mas 79-1, 07760 Ciutadella de Menorca, Illes Balears,
seeds could also be identified from material obtained in
Spain
3
Institut Mediterrani d’Estudis Avançats (CSIC-UIB), Ctra. Vallde-
a cave; the material here consists only of several frag-
mossa km. 7’5, 07122 Palma de Mallorca, Illes Balears, Spain
ments and one entire seed.
205

Material and methods
The material from Menorca described in this paper
has been obtained from a karstic site in Punta Nati (Ciu-
tadella), at the northwest of the island (deposit 11). Two
other deposits containing fossil seeds are known in the
island (deposit 28 of Racó des Pi and na Macaret; see
Fig. 1). The fossil seeds appear associated to different
elements of the so-called “the giant leporid fauna”, con-
sidered from the Pliocene. This fauna consists in a
highly peculiar vertebrate assemblage consisting in two
terrestrial mammals (an undescribed giant leporid and a
giant dormouse, Muscardinus cyclopeus), a bat (Rhino-
lophus
cf. grivensis), a giant tortoise (Cheirogaster
Figure 1. Map of Menorca showing the localities where fossil seeds
have been found and the only extant population of Cneorum
gymnesica), a lizard (Podarcis sp.), an undetermined
tricoccon in the island. 1. Punta Nati. 2. Racó des Pi. 3. Na
gecko, several snakes (Vipera natiensis, Vipera sp. and
Macaret. 4. Cala Mesquida.
an undetermined colubrid), an amphisbaenid (Blanus
Figura 1. Mapa de Menorca amb les localitats on s’han trobat llavors
sp.), a discoglossid (Latonia sp.) and different birds
fossils i la localitat on s’ha trobat l’única població vivent de
(such as Pterodromoides minoricensis, Scolopax car-
Cneorum tricoccon de l’illa. 1. Punta Nati. 2. Racó des Pi. 3.
mesinae, Camusia quintanai, Tyto balearica, Athene sp.
Na Macaret. 4. Cala Mesquida.
and Corvus sp.). This fauna was obtained after acid
acetic treatment of the blocks obtained in the deposits,
but the fossil seeds were obtained through physical
deposited in the private collection of Josep Quintana.
methods.
One of the objectives of the study was to compare
The fossil seeds from Eivissa were collected in the
the wall structure and the dimensions of C. triccoccon
cave of Ca na Reia, located in Puig des Guixer (Santa
seeds between fossil and current material. We examined
Eulària des Riu), at the northeast of the island and at an
the seed wall structure by means of a dissecting scope
altitude of 120 a.s.l. This cave consists of a number of
and measured seed diameter and length to the nearest
small cavities surrounding a collapsed large cavity.
0.01 mm of all seeds by means of a digital caliper.
Currently, the main cave consists in an opened hall of 13
x 7 x 6 m. The emergent sediment thickness is c. 3 m
and its length is c. 2 m. Detailed information on this cave
and its topography can be obtained from TRIAS (1982).
Systematic palaeontology
The obtained vertebrate fauna consists on two dormice
(Eliomys (Eivissia) canarreiensis and Eliomys sp.), bats,
a giant tortoise (Cheirogaster sp.), a lizard (Podarcis pit-
Family Cneoraceae Link, 1831
yusensis or its direct ancestor), and several species of
Genus Cneorum L., 1753
birds (among them, Puffinus nestori). This fauna has
Cneorum tricoccon L., 1753
been attributed to the Pliocene. The materials were
obtained without any special treatment of the sediment.
Catalan common name: Olivella or escanyacabres.
The materials from Eivissa are curated in the collec-
Material: 55 seeds obtained in Punta Nati locality 11
tion “Museu de la Naturalesa de les Illes Balears”, in
(Menorca). Several fragments of seeds obtained in cova
Palma de Mallorca, whereas those from Menorca are
de Ca na Reia (Eivissa). The species is also present in
Figure 2. Fossil seeds of Cneorum tricoccon collected at the karstic
Figura 2. Llavors de Cneorum tricoccon de Menorca col·lectades als
deposits from Menorca, after Quintana et al. (2004). Scale
jaciments càrstics de Menorca, segons Quintana et al. (2004).
bar, 5 mm.
Escala, 5 mm.
206

the deposit of Racó des Pi (south of Ciutadella; Quinta-
than 1 mm- that sometimes cover the fossil seeds. Any-
na 1198a; Quintana et al., 2004) and in na Macaret
way, the size of fossil seeds fall within the range (6.8-
(Quintana, 1998b). See Figs. 2 and 3.
2.72 mm in length and 1.92-5.51 mm in diameter; n=
Comments: Fossil seeds of C. tricoccon measure,
5781 seeds; Traveset, unpublished data) of those found
on average, 5.44 ± 0.68 mm and 4.71 ± 0.62 mm (x± sd;
in other populations of C. tricoccon in the Balearic
n= 55) in length and diameter, respectively. Both mea-
Islands and are known to vary from year to year depen-
sures were found to be somewhat higher than those
ding upon environmental conditions (RIERA et al.,
obtained from seeds collected at Cala Mesquida in the
2002).
fruiting season of 2002 (4.43 ± 0.50 mm and 4.00 ± 0.52
Finally, some Cneorum triccoccon seed fragments
mm, respectively, n = 414; Fig. 3A, B), the differences
were identified from Cova de Ca na Reia (e.g., Fig. 3C)
being highly significant (one-way ANOVAs, after loga-
together with an entire seed of Pistacia lentiscus.
rithmically transforming both parameters:
F1,467=150.38 and F1,467=71.83, respectively,
P<0.001). Such difference in size, however, might partly
Family Rosaceae Juss., 1789
be due to the layer of calcium carbonate -often thicker
Genus Crataegus L., 1753
Crataegus monogyna Jacq., 1775
Catalan common name: Cirerer de pastor, arç blanc
Material: A complete seed obtained in Punta Nati
locality 11 (Calas Pou). See Fig. 4.
Comments: Crataegus monogyna is a tree with irre-
gular leaves that currently spreads over humid areas in
the Balearic Islands.
The size of the Crataegus monogyna fossil seed is
6.37 mm in length and 4.71 mm in diameter, which also
falls within the range of those collected nowadays
(mean ± sd: 6.62 ± 0.82 mm; range: 5.1-8.3 mm for
seed length, and 4.89 ± 0.65, 3.86-6.12 for seed dia-
meter; n= 50 seeds).
Family Anacardiaceae Lindl., 1830
Genus Pistacia
Pistacia lentiscus
L., 1753
Catalan common name: Mata.
Material: One seed, collected in Cova de Ca na
Reia, Eivissa.
Comments: The entire seed of Pistacia lentiscus
(Anacardiaceae) is of a similar size to those collected
nowadays (Fig. 5).
Discussion
The best sample of fossil seeds was obtained from
a deposit of the North of Ciutadella. A total of 68 seeds
were gathered at Punta Nati locality 11 (Calas Pou). The
large majority of them (55; 81%) has been classified as
Cneorum tricoccon L. (Cneoraceae); they are very well
preserved and the characteristic seed shape and coat
structure facilitates their identification (see Figs. 2-3).
The rest of specimens cannot be attributed with cer-
tainty to any species except for one seed that we iden-
tify as Crataegus monogyna (Rosaceae) (Fig. 4). The
unidentified specimens from Calas Pou probably
Figure 3. Actual (A: Cala Mesquida) and fossil seeds (B: Punta Nati) of
Cneorum tricoccon from Menorca and Eivissa (C: Cova de Ca
belong, mainly or totally, to Cneorum tricoccon.
na Reia). Scale bar, 1 mm.
Cneorum tricoccon is a perennial shrub, often shor-
ter than 1 m, which belongs to a Mediterranean flora
Figura 3. Llavors actual (A: Cala Mesquida) i fòssil (B: Punta Nati) de
evolved during the early Tertiary under tropical condi-
Cneorum tricoccon de Menorca i d’ Eivissa (C: Cova de Ca na
Reia). Escala, 1 mm.

tions (RAVEN, 1973) and it thus represents a relict spe-
207

Figure 4. Actual (A) and fossil (B) seeds of Crataegus monogyna.
Figure 5. Actual (A) and fossil (B) seeds of Pistacia lentiscus from
Scale bar, 1 mm.
Eivissa. Scale bar, 1 mm.
Figura 4. Llavors de Crataegus monogyna de Menorca: (A) actual, i (B)
Figura 5. Llavors de Pistacia lentiscus d’ Eivissa: (A) actual i (B) fòssil.
fòssil. Escala, 1 mm.
Escala, 1 mm.
cies. It has a disjunct distribution in the western Medite-
The fossil seeds of C. tricoccon, obtained in
rranean basin. It is common in the Balearic Islands,
palaeontological deposits from Menorca (QUINTANA,
having been recorded on 14 islands and islets (Traveset
1998; QUINTANA et al., 2004, and the present study)
& Riera, unpubl. data), and has small populations in the
and located in sites where the plant is not currently pre-
southeastern Iberian Peninsula, southeastern France,
sent, are the only evidence that demonstrates the more
southwestern Sardinia and northwestern Italy (BOLÓS,
widespread distribution that this species had in this
1958; and Traveset, unpub. data). It usually inhabits
island. The shrinking of the populations of this species
coastal maquis or shrublands, usually in calcareous soils.
can be related to the disappearance of its main native
In the Balearics, the species has also colonized the inte-
disperser from Menorca, the endemic lizard Podarcis lil-
rior of the large islands; in Mallorca is often found in the
fordi (Lacertidae), as it seems to have happened with
understory of oak forests (Quercus ilex L.) up to an alti-
other plant species such as the endemic and threatened
tude of around 1000 m a.s.l. (TRAVESET, 1995, 2002).
shrub Daphne rodriguezii (TRAVESET & RIERA, 2005).
The fruit consists of three cocci that are easily separated
According to the fossil record, P. lilfordi began vanishing
from each other, and each coccus contains two one-see-
from the larger Balearic Islands, Mallorca and Menorca,
ded compartments. Information on its reproductive bio-
about c. 2000 years ago, presumably after the human
logy can be obtained in TRAVESET (1995 a,b).
introductions of predators such as weasels (Mustela
Crataegus monogyna is rather common both in
nivalis) (see ALCOVER, 1989; PLEGUEZUELOS et al.,
Menorca and Mallorca (BONAFÉ, 1979), and thus the
2002 and references therein). This lizard persists only in
finding of the fossil seed of this species is not unexpec-
the small islands around the two larger ones and it is an
ted. C. monogyna is currently distributed throughout
important seed disperser for a number of plant species
Menorca, mostly in the central part (Pere Fraga, pers.
(PÉREZ-MELLADO & TRAVESET, 1999).
com.) and is also abundant at Serra de Tramuntana in
In Mallorca, pine martens (Martes martes) appear
Mallorca. It is usually found in shady habitats, often near
to play an important role at present for the distribution of
temporary streams. Its fruits are consumed usually by
C. tricoccon as they consume large quantities of fruits
birds (Traveset, pers. obs.), although pine martens Mar-
and effectively disperse its seeds in that island (RIERA
tes martes probably contribute also to the dispersal of
et al., 2002; TRAVESET, 2002). In Menorca, in contrast,
this plant.
pine martens are not as frugivorous as in Mallorca
208

(CLEVENGER, 1993) and most fruits of C. tricoccon
BOUR, R. (1985): Una nova tortuga terrestre del Pleistocè d’Eivissa: la
tortuga de la cova de Ca na Reia. Endins, 10-11: 57-61.
remain undispersed (RIERA et al., 2002).
BOURROUILH, R. (1973): Stratigraphie, sédimentologie et tectonique
In the Pityusic Islands, the healthiest populations of
de l’ile de Minorque et du Nord-Est de Majorque (Baléares). La
C. tricoccon are found in Formentera and in the islets
terminaison nord-orientale des Cordilleres Bétiques en Médite-
south of Eivissa (TRAVESET & RIERA, unpub. data),
rrannée Occidentale. These. Paris. 822 pp.
BOVER, P. (2004): Noves aportacions al coneixement del gènere Myo-
where they have a strong interaction with the other
tragus Bate 1909 (Artiodactyla, Caprinae) de les Illes Balears. PhD
endemic lizard species of the Balearics, P. pityusensis.
Thesis, Universitat de les Illes Balears, 469 p.
The abundance of lizards in the larger island (Eivissa)
CLEVENGER, A .P. (1993): Spring and summer food habits and habitat
use of the European pine marten (Martes martes) on the island of
appears to have decreased in the last years (Riera,
Minorca, Spain. Journal of Zoology, London, 229: 153-161.
pers. com.) which may well have affected the popula-
COLOM, G. (1982): Notas paleontológicas y estratigráficas referents a
tions of this plant in the island.
Mallorca e Ibiza. Bolletí de la Societat d’Història Natural de les
Balears,
26: 195-206.
The Pistacia lentiscus seed found in Cova de Ca
DEPAPE, G. & FALLOT, P. (1928): Les gisements du Burdigalien a plan-
na Reia is not surprising considering that this species is
tes de Majorque. Ann. Soc. Géol. Nord, 53 : 5-19.
also very ancient, from the early Tertiary (RAVEN,
HABLY, L. & FERNÁNDEZ MARRON, M.T. (1998): A comparison of the
Oligocene floras of the Tethyan and Central-Paratethyan areas on
1973), and quite common in many habitats in all the
the basis of Spanish and Hungarian macroflora. Tertiary Research,
Balearics (BONAFÉ, 1979).
18: 67-76.
Finally, the presence of Cneorum tricoccon in the
KÖHLER, M. & MOYÀ-SOLÀ, S. (2004): Reduction of brain and sense
Pliocene deposits of the Balearic Islands confirms the
organs in the fossil insular bovid Myotragus. Brain Behavior and
Evolution
, 63: 125-140.
antiquity of the presence of this species in the Balearics.
LLOMPART, C.; OBRADOR, A., & ROSELL, J. (1979): Geologia de
In our knowledge, these findings represent the first fos-
Menorca. In VIDAL, J.M. (coord.), Geografia física. Enciclopèdia
sil documentation of the species, and it is highly consis-
de Menorca, I: 5-83.
PÉREZ-MELLADO, V., and TRAVESET, A. (1999): Interactions betwe-
tent with the interpretation of its belonging to the subtro-
en Mediterranean lizards and plants. Natura Croatica, 8: 275-285.
pical flora that was once spread in the Balearics, before
PLEGUEZUELOS, J.M.; MÁRQUEZ, R.& LIZANA, M. (2002): Atlas y
the start of the glaciations.
Libro Rojo de los Anfibios y Reptiles de España. Dirección Gene-
ral de Conservación de la Naturaleza-Asociación Herpetológica
Española. Madrid.
QUINTANA, J. (1998a): Aproximación a los yacimientos de vertebrados
del Mio-Pleistoceno de Menorca. Bolletí de la Societat d’Història
Natural de les Balears,
41 : 101-118.
Acknowledgements
QUINTANA, J. (1998b): Presencia de Trochoidea frater (Dohrn and
Heynemann, 1862) (Gastropoda: Helicidae) en los depósitos cárs-
ticos de Menorca. Bolletí de la Societat d’Història Natural de les
Balears
, 41: 49-56.
This paper is included in our Research Projects
QUINTANA, J. (2005): Estudio morfológico y funcional de Nuralagus
BOS2001-0610 and CGL2004-04612/BTE of the Direc-
rex (Mammlia, Lagomorpha, Leporidae). PhD Thesis, Universitat
ción General de Investigación, Ministerio de Educación
Autònoma de Barcelona, 360 + 142 pages.
y Ciencia (Madrid). We thank Javier Tomás for his assis-
QUINTANA, J.; BOVER, P. & ALCOVER, J.A. (2004): Els vertebrats
terrestres fòssils del Migjorn. In FORNÓS, J.J.; OBRADOR, A. &
tance with photographs of seeds under the scope and
ROSSELLÓ, V.M. (eds.), Història Natural del Migjorn de Menorca:
the Jardí Botànic de Sóller for letting us consult their
el medi físic i l’influx humà. Monografies de la Societat d’Història
seed collection.
Natural de les Balears, 11: 291-302.
RAMOS, E. & ÁLVAREZ, C. (1989-90): Paleoflora del Oligoceno de
Mallorca. Bolletí de la Societat d’Història Natural de les Balears
31: 141-158.
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