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International Journal of Paleopathology
journal homepage: www.elsevier.com/locate/ijpp
Paradise lost: Evidence for a devastating metabolic bone disease in an
insular Pleistocene deer
George A. Lyras
a
, Aggeliki Giannakopoulou
b,
⁎
, Theodoros Lillis
c
, Alexandra A.E. van der Geer
d
a
Faculty of Geology and Geoenvironment, National and Kapodistrian University of Athens, Athens, Greece
b
Laboratory of Anatomy, Histology and Embryology, School of Veterinary Medicine, Faculty of Health Sciences, Aristotle University of Thessaloniki, Thessaloniki, Greece
c
Department of Oral Surgery, Implantology and Radiology, School of Dentistry, Faculty of Health Sciences, Aristotle University of Thessaloniki, Thessaloniki, Greece
d
Naturalis Biodiversity Center, Leiden, the Netherlands
ARTICLE INFO
Keywords:
Candiacervus
Mavromouri
Palaeohistology
Nutritional secondary hyperparathyroidism
Osteophagia
Bone remodeling
ABSTRACT
Purpose: This communication reports skeletal pathology in a Pleistocene endemic deer from the Mavromouri
caves of Crete.
Materials: 287 bones and bone fragments from Mavromouri caves are compared to 2986 bones from Liko Cave.
Methods: Bones were evaluated macroscopically, and measurements were made of morphometric characteristics
of limb long bones. Representative bone specimens were examined radiographically and histologically.
Results: Macroscopic hallmarks were loss of bone mass and increased porosity. The long bones were brittle, some
of them having thin cortices, and others reduction of medullary cavities that contain dense Haversian tissue. The
flat bones were spongy and fragile. Erosions of the metaphyses and articular surfaces were noted. Histological
findings included: sub-periosteal resorption; loss of lamellar bone; enlargement of vascular canals; and re-
modeling of cortical bone. Two types of fibrous osteodystrophy were recognized in skeletal remains, subostotic
and hyperostotic.
Conclusions: The deer of Mavromouri caves were affected by severe metabolic bone disease, likely nutritional
secondary hyperparathyroidism. We hypothesize a multifactorial cause, including overgrazing, flora senescence,
soil mineral deficiencies, and a prolonged period of climate extremes, degrading the Cretan deer habitat.
Value: This is the first evidence of a metabolic bone disease causing this level of destructive pathology in an
insular fossil deer.
Limitations: The lack of absolute chronometric dates for the site limits potential linking with the prevailing
environmental conditions.
Suggestions for further research: Investigation of similar skeletal pathologies at other islands or isolated habitats is
advised.
1. Introduction
Endemic ruminants are known from several Eurasian palaeo-islands.
There are many studies of the phylogeny, anatomy, and life history of
ancient insular ruminants (van der Geer et al., 2010), but works on their
pathology are few (Waldren, 1999; Jordana and Kohler, 2011; Palombo
and Zedda, 2016). Paleoepizootiological approaches considering pos-
sible reasons for their extinction also are rather rare (De Souza et al.,
2003). Investigations of fossil remains and their pathology contribute to
our understanding of the island ecosystems and perhaps of eventual
species demise (Rothschild and Martin, 1993).
Crete was an island throughout the entire Pleistocene, without
previous land connection; thus, mammals could reach it only by aquatic
dispersal (van der Geer et al., 2015). The few mammalian species that
successfully colonized Crete in the Middle and Late Pleistocene are
limited to a dwarf elephant (Palaeoloxodon creutzburgi), a large mouse
(Mus minotaurus), a shrew (Crocidura zimmermanni), an otter (Lutrogale
cretensis), and the Cretan deer (Candiacervus, 8 species) (van der Geer
et al., 2010; van der Geer, 2018). All of those species now are extinct,
except for the shrew.
Cretan deer likely inhabited the entire island but fossils are re-
stricted to coastal caves around the island, except for the southern coast
and a few inland sites (Iliopoulos et al., 2010). The largest collection of
Cretan deer fossils derives from a single site, Liko Cave near the village
Likotinara, and consists of more than 6000 specimens. Other important
collections have been retrieved from coastal caves in the Rethymnon
https://doi.org/10.1016/j.ijpp.2018.12.003
Received 6 May 2018; Received in revised form 29 November 2018; Accepted 11 December 2018
⁎
Corresponding author. Present address: Department of Hematology, G. Papanikolaou General Hospital, Thessaloniki, Greece.
E-mail address: agiann@auth.gr (A. Giannakopoulou).
International Journal of Paleopathology 24 (2019) 213–226
1879-9817/ © 2018 Elsevier Inc. All rights reserved.
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