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M. Masseti: Homogenisation and the loss of biodiversity of mammals of the Mediterranean islands 175
Figure 8 Ideal reconstruction of the elephant of Falconer, Ele-
phas falconeri Busk 1867, which was only 0.90 m tall at the
withers, thus representing the smallest elephant that ever lived ten and Sondaar 1972), being about the size of a pig
(courtesy of the Muse´ um National d’Histoire Naturelle, Paris). (Sondaar 1991) (Figure 9). Furthermore, the odontology
native biocenoses, and introducing many new species.
Virtually no ecosystems have been left untouched. Few
other regions of our planet have been maltreated by man
more than the Mediterranean, to the extent that we can
now have only the vaguest impression of what the
ancient natural riches, vegetable lushness and primaeval
faunal luxuriance of the region may have been. Much of
the environment of the Mediterranean islands is by now
reduced to little more than a mineral skeleton. In the
Upper Pleistocene, when the existing botanical and fau-
nal assemblages were already defined, the majority of the
Mediterranean islands must still have been cloaked in the
densely tangled vegetation that covered most of these
environments before any human explorer set foot on
them. Oak jungles consisted of downy oaks, Quercus
pubescens Willd., evergreen oaks, Q. ilex L. 1753, Ker-
mes oaks, Q. coccifera L. 1753, and of endemic species,
such as the golden oak of Cyprus, Q. alnifolia Poech
1842, (cf. Haslam et al. 1977, Rackham 1978, 1990,
Turner 1978, Meikle 1977, Turland et al. 1993, Rackham
and Moody 1996, Ntinou 2002, Masseti 2008). Not to
mention the dense undergrowth. The ‘‘true’’ islands of the
Mediterranean were inhabited by extraordinary creatures
which might have proved to be intriguing aberrations
from the most simple rules of evolution. These practically
impenetrable vegetable formations must have been
home to an entire fauna that was particularly adapted to
living, moving, feeding, resting and hunting each other in
such an environment. This is still the case, for example,
in certain relic fragments of the rain forests of western
Africa, where peculiar biological elements, such as the
last pygmy hippopotami, Hexaprotodon liberiensis Mor-
ton 1849, still survive. This is a species dispersed at
present only in western Africa from Sierra Leone to the
Coˆ te d’Ivoire (cf. Grubb 2005). Its main habitat consists
of forested watercourses where it shelters by day in
ponds, rivers and swamps. At night, it follows tunnel-like
paths through dense riverine vegetation to graze in
glades, or along grassy trails (Kingdon 2004). Recently,
pygmy hippos have also been ‘‘rediscovered’’ in Liberia,
in the Sapo National Park. It cannot be excluded that
some of the dwarf hippopotami of Sicily, Malta, Crete and
Cyprus might have lived in similar environmental condi-
tions too. The Cypriot dwarf hippo was even smaller and
more slender than the extant Liberian hippo (Boekscho-
and morphology of the Cypriot species suggest a mode
of living similar to that of leaf-eating pigs and it may have
been an animal that was better adapted to walking than
to swimming (Boeckschoten and Sondaar 1972). The
dwarf endemic elephant which characterised the Late
Pleistocene/Holocene faunal horizons of the small island
of Tilos, in the Eastern Aegean sea (Greece), has been
described as a very agile animal well adapted to the
island environment, capable of moving through rough
terrain (Theodorou et al. 2007b).
It seems plausible that the reduced body size of the
large mammals was one of the best adaptive responses
to this type of environment. Analogous examples can be
found in the populations of other large herbivores that
still populate the primeval forest regions of Africa. Dwarf
populations of African buffaloes, for example, taxonom-
ically defined as Syncerus caffer nanus Boddaert 1785
(Basilio 1962, Eisentraut 1973, cf. Grubb 2005), and also
known as forest, red or dwarf buffaloes, are regarded as
still occurring in what remains of the African primeval for-
ests, from Gambia to southern Zaire and northern Ango-
la, and eastwards to lakes Edward and Kivu and
south-western Ethiopia (Halthenorth and Diller 1977,
Yalden et al. 1984).
Elephants of reduced body dimensions are still reputed
to inhabit the rain forests of west-central Africa too (Basi-
lio 1962, Roeder 1970, 1975, Halthenorth and Diller 1977,
Kingdon 2004, Shoshani 2005). However, the existence
of a species of African pygmy elephant is not generally
recognised. In this regard, Halthenorth and Diller (1977)
observed that they are often believed to be small exem-
plars of the African forest elephant, Loxodonta cyclotis
Matschie 1900. The latter form was until recently consid-
ered a subspecies of the African bush elephant, Loxo-
donta africana Blumenbach 1797, but DNA testing has
now shown that there may possibly be two extant ele-
phant species in Africa. In fact, the forest elephant has
recently been shown to be genetically distinct from the
Figure 9 Artist’s reconstruction of the extinct pygmy hippo-
potamus Phanourios minor Desmarest 1822, of Late Pleistocene
Cyprus, adapted from the specimen in the London Natural His-
tory Museum and compared in size with the extant Hippopota-
mus amphibius L. 1758 (Drawing by Alessandro Mangione).