Microfauna from historical sites in the Aegean
An assessment of the excavated evidence and issues of small mammal synanthropy and commensalism
p. 157-175
Résumés
This paper summarises the microfaunal evidence from Aegean sites of the 1st millennium BC. Microfaunal skeletal material of rodents (mice, rats, hamsters, etc.), insectivores (shrews, moles, bats, etc.), reptiles (small snakes and lizards), amphibians (frogs, toads, salamanders) and small-sized birds has not systematically been collected from historical sites; however, the limited data permit the reconstruction of open landscapes in alternation with maquis vegetation and Mediterranean woodland in the areas around the sites. Furthermore, the limited evidence permits the identification of commensal species (house mice, rats) and allows a short discussion of the biogeography of those genera. Finally, reptile skeletal material from a 1st c. BC sanctuary of Poseidon is discussed in terms of ritual use and targeted deposition.
Cet article fait le point sur la microfaune dans les sites de l’Égée au Ier millénaire av. J.-C. Les restes osseux de la microfaune des rongeurs (souris, rats, hamsters, etc.), insectivores (musaraignes, taupes, chauves-souris, etc.), reptiles (petits serpents et lézards), amphibiens (grenouilles, crapauds, salamandres) et oiseaux de petite taille n’ont pas été collectés de manière systématique sur les sites historiques ; les données, limitées, permettent cependant de reconstituer des paysages ouverts alternant avec des maquis et des forêts méditerranéennes dans les secteurs proches des sites. En outre, les sources permettent d’identifier des espèces commensales (souris domestiques, rats) et d’esquisser une discussion sur la biogéographie de ces espèces. Enfin, des restes osseux de reptiles d’un sanctuaire de Poséidon, au Ier siècle av. J.-C., sont interprétés comme issus d’usages rituels et d’un dépôt intentionnel.
Entrées d’index
Mots-clés : micromammifères, reptiles, commensalisme, habitat, rituel
Keywords : micromammals, reptiles, commensalism, habitat, ritual
Texte intégral
Introduction
1Microfauna is a complex of small-sized vertebrates of different taxa, either terrestrial mammals, reptiles, amphibians or small birds. The respective mammals are of small or minuscule size and they usually weigh between some grams and five kilograms; the commonest taxa found in archaeological sites are rodents and insectivores.1 Targeted soil sampling and washing techniques, such as water sieving and flotation, are necessary to recover microfaunal elements, as they are not visible during excavation by naked eye. Small mammals are a necessary tool to complement the archaeological and historical knowledge for natural habitat shifts and biogeographic changes facilitated by humans.2
Biochronology, biogeography, habitat and commensalism: the usefulness of micromammal studies in archaeology
2Micromammal species are a common research tool of vertebrate palaeontologists and Palaeolithic archaeologists for the reconstruction of paleoenvironments, as well as for the relative dating of sites via biostratigraphic evidence. The presence and the association of fossil small mammals in the sediments is called biostratigraphy,3 which provides the relative chronology of the geological or archaeological layers. Palaeontologists date the first and the last appearance of a genus or species in the stratigraphy and pay special attention to the evolutionary stage of a species or genus in a given site: all these information comprise the biostratigraphy.4 The biochronology suggested by the biostratigraphic data is then associated with absolute dates, provided by methods such as palaeomagnetism,5 when available. The biostratigraphic system is also applied in the zooarchaeology of the Pleistocene, and especially of the Lower Palaeolithic period.6 Furthermore, changes in climate during the Palaeolithic period are strongly expressed in the rodent fauna.7 Micromammals, rodents in particular, are useful in environmental reconstruction, since they have narrow microhabitat requirements and can be used as environmental indicators.8
3On the contrary, Holocene micromammal research does not offer relative chronologies, because the respective periods are very short compared to the Pleistocene ones, in order for the small mammal genera to have naturally changed. However, the usefulness of small mammals in providing palaeoecological and environmental indications about habitat shifts is important for Holocene studies too. Furthermore, Holocene small mammals provide biogeographic information regarding the distribution of specific species around the earth, either due to natural or human agency.9 A useful example is that of Corsica: natural habitat shifts are indicated by the gradual extinction of four Pleistocene endemic small mammals due to competition with eight species gradually imported by humans as early as the Middle Neolithic period; they were transported as stowaway boat cargoes in a period lasting almost seven millennia (mid-5th millennium BC to ~ 1300 AD) and due to the subsequent modification of the landscape by humans for agricultural practices.10
4Certain small mammals found in the anthropogenic habitat can betray synanthropic or commensal behaviour under specific circumstances in human history. The term synanthropic refers to the close spatial association with people, whereas the term commensal, deriving from the Latin cum (together) and mensa (table),11 denotes the trophic benefits from this association12 and implies that commensals live in the same house with humans.13 Synanthropic animals do not necessarily rely on humans for survival, despite cohabitation with them in anthropogenic environments.14 Usually the commensal relation between humans and mice starts as a synanthropic one.15 Commensalism is an intense but also indirect relationship of coexistence, where the passive host (human) does not seek direct benefit from the active commensal (mouse), while the latter seeks shelter/safety and food in the human niche.16 When both partners are benefitted the relation is called mutualism,17 whereas when the host is harmed, the relation is called parasitism.18 Commensals can harm humans while infesting their homes and cellars, because they carry pathogens either as insect parasites or in their excrements.19 Since such kinds of relationships presuppose that partners share their habitat or home, house mice and rats are the perfect commensals and their ecological preferences allow them to be indirectly translocated by humans in areas where they are not naturally distributed.20 A good example is the house mouse, which is the commonest mammal around the globe,21 the dispersal of which is a result of human activity.22 Due to this complex relationship between humans and mice, the term synanthropy can be used as an alternative to commensalism.23
The house mouse and the black rat: human-mediated dispersals and domestication
5The global spread of the house mouse (Mus musculus) occurred due to its synanthropic character and its adaptability to the human niche. It originates in Central Asia and the Indopakistani subcontinent24 and it was naturally dispersed in the Middle East by the Upper Pleistocene.25 It was as early as the Natufian period (13000-9500 BC), when it colonised the human niche taking advantage of the human waste and the lack of predators in the earliest sedentary settlements of Israel and Syria.26 The development of agriculture and the storage of cereals during the Pre-Pottery Neolithic A, B and C periods (10000-7000 BC, henceforth PPN A, B or C) brought the house mouse to the Levantine settlements, from where it was spread in Cyprus and Anatolia along with the migrating human groups.27 There are direct C14 dates of mouse bones found in PPN A Ganz Dareh in Iran (8153-7483 cal BC) and PPN B Çatal Höyük (8248-4583 cal BC) and Cafer Höyük in Anatolia (8829-7169 cal BC). The oldest house mouse of Europe comes from the Chalcolithic settlements Buçsani La Pod Tell (Romania) and Vinča-Belo Brdo (Serbia), where it arrived through the Caucasus and the Black Sea or the Pontic Steppes. The Buçsani mouse was C14 dated to 4619-4464 cal BC.28 The intensification of commercial contacts between the peoples of the Mediterranean brought it to the Early Bronze Age layers of Akrotiri on Thera, directly dated to 2502-1913 cal BC.29 The Minoan commerce was also responsible for further distribution of the species in Neopalatial settlements on east and west Crete.30
6The black rat (Rattus rattus) is another typical commensal rodent and endemic of southeast Asia,31 the dispersal of which is also associated with humans, and especially with urbanisation. Like the house mouse, it had a natural expansion in the Near East by the Early Holocene, since it is found in the Natufian layers of Abu Usba and Hayonim caves in Israel, as well as in the PPN layers of Nahal Sefunim rock shelter and Sefunim cave.32 A lack, so far, of Rattus remains from open air PPN sites pinpoints to either a pause in the early rat commensalism or indicates the natural origin of the rat remains found in Natufian caves.33 The black rat (Rattus cf. rattus) reappeared in Pottery Neolithic Wadi Shu‘eib in Jordan.34 However, the true cradle of its commensalism is considered to be the Indus valley.35 From there it was distributed to the Middle East by merchants, due to the commercial contacts of the Mesopotamian cultures with the Harappan civilisations, since its remains are found in sites of the Tigris and Euphrates basin dated between 2500 and 1500 BC.36 A recent paleogenomic study of black rat remains supports such an early terrestrial expansion of the species to the Mediterranean via southwest Asia during the 2nd millennium BC.37
7The earliest certain black rat finds outside Asia are of dubious stratigraphic security: Neolithic presence of black rat in Su Guanu in Sardinia dated about 3500 BC38 and a Bronze Age rat from Cerro del Real, Spain39 could be a result of post-depositional contamination.40 Secure are considered the rat bones from Korucutepe in Anatolia, dated between 1400-1200 BC.41 Occasional rat isolated molars or isolated postcranial elements have been found in Aegean Bronze Age sites too: in Early Bronze Age Chania town layers (Crete), in Late Bronze Age Mochlos (Crete)42 and Katafygadi cave (Kythera island) layers.43 However they are considered post-depositional intrusions, due to their preservation state and the scarcity in their presence. Similarly, there is mention of black rat remains from 5th-4th c. BC sites in Poland,44 for which there is also doubt in terms of stratigraphic safety.45 More stratigraphically secure specimens from 2nd millennium BC sites are necessary to support an early permanent establishment.46
8The real dispersal of the species took place after the Roman conquest of the Mediterranean, when it was accidentally dispersed by boats. Armitage et al.47 consider its introduction to Europe a result of marine transport of this species from India to Egypt: during the Ptolemaic period exotic products were brought to the ports of the Red Sea inside boat cargoes. In this way black rats infested Egyptian towns, since their skeletons are found inside the guts of mummified birds of prey.48 Once Romans conquered Egypt, they maintained the commercial networks with India and thus more black rats were transported by chance under the decks of commercial boats. A black rat skeleton came to light in the stomach of a mummified cat dated to the 2nd-1st c. BC at Quseir el Qadim in Egypt, the port of Myos Hormos.49 At the time, Quseir el Qadim flourished as a port for the import and redistribution centre for spices from India. Subsequently, the black rats were dispersed around the Mediterranean, wherever the Roman rule was in effect, because Egypt was the main granary of the Roman Empire. Inside grain cargoes, the black rat arrived in several sites of the Roman period in the west Mediterranean: Pompei (Italy, end of 2nd c. BC), Ordona (Italy), Monte di Tuda (Corsica, 393-151 BC), Carthago and Zembra-Abri du Casino (Tunisia, Roman period), Menorca (2nd c. BC), Quinta do Marim (Portugal, 250-350 AD).50 It has also been found in the Roman layers of several Aegean sites, as discussed below.
Native micromammals of the Aegean and their palaeoenvironmental indications
9Skeletal elements of microfauna have been recovered from various Aegean sites dating between the 11th and the 1st c. BC (fig. 1). In most of them, the microfaunal material was collected randomly at the trench or through dry sieving, thus consisting of large-sized genera and of few specimens. Wherever soil flotation or water sieving was applied, the species richness was higher and the specimens more numerous, like in the 11th-10th c. BC Nichoria, Late Hellenistic/Early Roman Pyrgouthi or in the Roman sanctuary of Poseidon at Poros (3rd-1st c. BC), providing evidence for the palaeoecology of the region or for ritual practices.51 This patchy available data, however, do not permit a coherent palaeoenvironmental reconstruction of the small mammal biodiversity of the different regions through time. For consistency reasons, data providing environmental information from native microfauna will be presented separately from evidence for imported commensal/synanthropic species.
Fig. 1. Historical sites mentioned in the text and tables 1-2.

1: Nichoria, 2: Kastanas, 3: Mesimeriani Toumba, 4: Kabirion (Thebes), 5: Altar of Aphrodite Ourania (Agora of Athens), 6: Corinth, 7: Mytilene (polis and Acropolis), 8: Pyrgouthi, 9: Poseidon Sanctuary (Kalaureia, Poros), 10: Kommos Temple C, 11: Kavoussi, 12: Eleutherna, 13: Knossos. Empty circles: Late Dark Ages; filled circles: Geometric period; empty squares: Archaic period; filled squares: Classical period; empty triangles: Hellenistic period; filled triangles: Roman period.
10In sites where systematic soil screening was not practiced, large-sized genera are usually found, which belong to the local/endemic fauna of continental Greece: squirrel (Sciurus vulgaris), ground squirrel (Spermophilus citellus), hedgehog (Erinaceus europaeus), lesser mole-rat (Nanospalax leucodon) (table 1). All these genera are naturally distributed around Greece today and pinpoint to an open vegetation area, grassland or sparse woodland.52 The squirrel and ground squirrel were recovered from Geometric layers at Kastanas (Central Macedonia); the squirrel mandible bears cut marks, which betray skinning efforts for the animal’s fur.53 The hedgehog was identified at Kastanas Dark Age layers,54 in Archaic/Late Classical and Roman layers at Mytilene Acropolis55 and at Eleutherna Hellenistic/Roman layers;56 its presence at Kastanas has been associated with exploitation of its spikes as needles or its meat as healing food.57 Four bones from one squirrel were recovered from the Geometric House at Mesimeriani Toumba58 (Thessaloniki). At the Kabirion of Thebes, one bone of Hellenistic date and two bones of Roman date were recovered, all belonging to two lesser mole-rats.59 The lesser mole-rat was also identified in Late Hellenistic/Early Roman layers at Pyrgouthi (Argolid, four bones).60 Finally, at the shrine of Glaukos at Knossos eight non-identified small mammal bones have been recovered.61
11Wherever soil flotation or sieving was practiced, smaller-sized micromammal species were recovered. The available data come from two settlements in the Peloponnese, a sanctuary on Poros island and one temple in Crete: Nichoria (Messenia),62 Pyrgouthi (Argolid),63 Poseidon sanctuary in Kalaureia (Poros island)64 and Kommos Temple C (Crete).65 The identified species indicate either an open landscape with maquis bushes or cultivated fields with bushes at their edges or Mediterranean woodland: shrew (Crocidura sp.), greater white-toothed shrew (C. russula), Etruscan shrew (Suncus etruscus) [pl. III, 18], edible dormouse (Glis glis) [pl. II, 11], common dormouse (Muscardinus avellanarius) [pl. II, 12], forest dormouse (Dryomys nitedula), vole (Microtus sp.), snow vole (Microtus nivalis), wood mouse (Apodemus sylvaticus), yellow-necked mouse (A. flavicollis), rock mouse (A. mystacinus) (table 1).
Table 1. Micromammal species from sites mentioned in the text. Numbers refer to actual bones (Numbers of Identified Specimens, NISP); MNI stands for Minimum Number of Individuals; x denotes presence in the layers.
Nichoria | Kastanas | Mesimeriani Toumba | Kabirion (Thebes) | Altar of Ourania Aphrodite | Roman Corinth | Mytilene (Acropolis and Polis) | Pyrgouthi, Argolid | Poseidon Sanctuary, Kalaureia | Kommos Temple C | Kavoussi | Eleutherna | Roman Knossos | |
Crocidura russula | 28 | ||||||||||||
Crocidura sp. | x | ||||||||||||
Suncus etruscus | 1 MNI (~ 325 BC) | ||||||||||||
Erinaceus concolor | 1 | x (Archaic/Late classical, Roman) | 1 | ||||||||||
Glis glis | 5 | ||||||||||||
Muscardinus avellanarius | 2 | ||||||||||||
Dryomys nitedula | 1 | ||||||||||||
Sciurus vulgaris | 1 | 4 (1 MNI) | |||||||||||
Spermophilus citellus | 1 | ||||||||||||
Chionomys nivalis | 11 | ||||||||||||
Microtus sp. | 1 | ||||||||||||
Muridae | x (Roman) | 7 | |||||||||||
Apodemus sylvaticus | 19 | 14 | |||||||||||
Apodemus flavicollis | 49 | ||||||||||||
Apodemus mystacinus | 32 | 4 (~165 BC) | 1 | ||||||||||
Acomys minous | 9 | ||||||||||||
Mus musculus | 20 | 157 (13 MNI) | 2 (1 MNI) | 5 (2 MNI) | 3 (50 BC-100 AD) | 71 | |||||||
Rattus sp. | 1 | 23 (50 BC-100 AD) | 1 | ||||||||||
Rattus rattus | x (Archaic/Late classical, Roman) | 2 | 6 (4 MNI) | ||||||||||
Nanospalax leucodon | 3 (1 Hellenistic, 2 Roman) | 4 (2 MNI) |
12The Nichoria microfaunal assemblage was found inside a pithos dated between the Late Dark Ages (1050-900 BC) and the Early Geometric period (900-850 BC) that functioned as a natural trap; the assemblage comprised more than 200 individuals, including amphibians and reptiles along with micromammals.66 The Nichoria micromammal species still exist in Peloponnese and Crete, apart from the snow vole (Microtus nivalis67), an alpine species, and the greater white-toothed shrew (C. russula), which are both currently not distributed in the Peloponnese.68 The deposition of Pyrgouthi and Kommos Temple bones is attributed to remnants of owl pellets, like those produced by Tyto alba. Owls fed on the synanthropic species that lived in the cultivated land around Pyrgouthi or Kommos and roosted in empty or infrequently used buildings, such as the Temple C.69
Commensal imported species from historical sites in the Aegean
13When it comes to commensal non-native micromammals, there is also scanty evidence: black rat and house mouse bones have been occasionally retrieved (table 1). The black rat (Rattus sp., R. rattus) has been identified at Kavoussi (Geometric period, Crete; two bones),70 Eleutherna (Hellenistic/Roman, Crete; six bones),71 the Acropolis of Mytilene (Archaic/Late Classical and Roman layers; no count),72 Pyrgouthi (Late Hellenistic/Early Roman; one bone), Knossos (Roman layers; one bone),73 Poseidon sanctuary at Kalaureia (Roman layers; 23 bones).74 The house mouse (Mus musculus) was identified in Late Dark Age (1050-900 BC)-Εarly Geometric layers (900-850 BC) at Nichoria (20 bones),75 in the Iron Age Temple C at Kommos (71 bones),76 in Late Hellenistic/Early Roman layers at Pyrgouthi (five bones),77 in a room of Roman Corinth (one individual)78 and in 1st c. BC layers at Poseidon sanctuary in Kalaureia (three bones).79 An assemblage of 13 house mice (157 bones) was recovered from the Classical period Altar of Aphrodite Ourania at the Agora of Athens,80 so far the largest house mouse assemblage from 1st millennium BC Greece. The presence of the house mouse in Attica and the Peloponnese reflects the establishment of the species in the Aegean since the Early Bronze Age.81 The rat presence in the Roman layers of the aforementioned sites is anticipated, because the species was distributed across the Mediterranean during the Roman times.82 The rat bones from Geometric Kavoussi are thus considered very early and might be intrusive; this might also be the case for the Archaic/Late Classical rat from the Mytilene Acropolis, where bioturbation might have mixed rat bones from Roman layers into earlier strata.
14An unexpected import was found in a Hellenistic layer of Temple C at Kommos (375 BC-160/170 AD): the spiny mouse (Acomys sp.) [pl. III, 16]. The genus Acomys is a native of west Asia and North Africa existing in the paleontological record of those regions. Today an endemic insular form, Acomys minous [pl. III, 17], is located on Crete, which evolved locally after multiple introductions of individuals originating from continental populations.83 Humans introduced it to Crete unintentionally, probably in commercial boat cargos.84 The Kommos find constitutes its earliest so far presence on the island.85 An endemic spiny mouse, Acomys nesiotes, is now living on Cyprus and it is also thought to have been accidentally imported by humans after the Bronze Age potentially from Crete.86 However, lack of prehistoric or ancient evidence of this species dates its earliest appearance on Cyprus during the last 1,000 years.87
Non-mammalian microfaunal evidence from historical sites in the Aegean
15Reptile or amphibian remains from sites spanning the 1st millennium BC are even scantier than the micromammal ones, thus not permitting coherent palaeoenvironmental observations (table 2). The assemblages from Nichoria and Poseidon sanctuary area D at Kalaureia are the most detailed ones. The Nichoria microfaunal assemblage included several amphibian and reptile elements belonging to: common toad (Bufo bufo spinosus), Greek frog (Rana graeca) [pl. IV, 38], lake frog (R. ridibunda), marsh frog (cf. Pelophylax sp.), green lizards (Lacertilia, L. viridis), gecko lizards (Gekkonidae) and various snakes88 (table 2), all native to continental Greece.
Table 2. Amphibian and reptile species from sites mentioned in the text. Numbers refer to actual bones (NISP), whereas “x” denotes presence in the layers.
Nichoria | Kaveirio (Thebes) | Altar of Ourania Aphrodite | Ancient Corinth | Mytilene (Acropolis and Polis) | Poseidon Sanctuary, Poros | |
Bufo bufo spinosus | 30 | |||||
Rana graeca | 1 | 6 | ||||
Rana ridibunda | 1 (Late Hell/Roman) | |||||
Pelophylax sp. | x | |||||
Anura | 1 | x | ||||
Lacertilia, L. viridis and geckos | 27 | x | ||||
Reptilia | ||||||
Ophidia | 3 | |||||
Hierophis gemonensis | x | |||||
Malpolon monspessulanus | x | |||||
Elaphe quatuorlineata | x | |||||
Zamenis longissima | x | |||||
Telescopus fallax | x | |||||
Vipera ammodytes | x | |||||
Natrix natrix | x | |||||
Natrix tessellata | x |
16The zooarchaeological studies of the sanctuary of Poseidon at Kalaureia reveal ritual practices that incorporate microvertebrate species on purpose. The systematic soil flotation yielded microfauna from all layers of the Sanctuary (750-700 BC through 50 BC-AD 100; table 1-2). The non-mammalian microfauna includes: lizards smaller in size than the Lacertidae family, snakes and marsh frogs (cf. Pelophylax sp.).89 In total, 2,398 various snake bones belonging to eight different species where recovered from all sanctuary layers: Balkan whip snake (Hierophis gemonensis) [pl. IV, 36], Montpellier snake (Malpolon monspessulanus), four-lined snake (Elaphe quatuorlineata), grass snake (Natrix natrix), dice snake (Natrix tesselata), nose-horned viper (Vipera ammodytes), Aesculapian snake (Zamenis longissima) [pl. IV, 37] and cat snake (Telescopus fallax). The identified viper is venomous, and the grass and dice snakes are fresh water snakes.90 The majority of the microfauna, and especially of the snake remains (1,544 bones) came from the fill of a Late Hellenistic/Early Roman cistern (50 BC-AD 100). The animal material from that cistern included also a donkey, a young horse, a bovine, pigs, caprines, dogs, fish, birds, eggshell and purple shells.91 Some of the snake ribs and vertebrae from the cistern were burnt (41 black bones) or recrystallised (37 white bones) or burnt/recrystallised (eight bones) indicating charring over fire as part of meals or of some ritual activity. Snake skull and jaw fragments did not bear any surface modification.92 Apart from one frog bone, the rest of the microfaunal remains (micromammals and lizards) did not indicate any burning evidence.93 This unusual assemblage was interpreted as ritual: snakes and lizards were conceived of chthonic character during classical times; snakes referenced the underworld, they were associated with Poseidon, Hekate and were the symbol and sacred animal of Asclepius.94 Gecko or other lizards were occasionally used for healing human eyes or were used in charms of sexual attraction or personal strength.95 Furthermore, all animals found in the cistern can be associated with divination or magic.96 All frogs, snakes and lizards identified in the cistern belong to the native modern Greek fauna and were not imported in the Sanctuary from outside Greece for religious purposes. However, it remains unknown whether they were captured in the immediate vicinity of the Sanctuary or if they were transported from further away.
Conclusions
17This paper summarized the evidence of microfauna from historical sites in the Aegean. A basic conclusion is the scarcity of the material, due to inadequate soil sampling and screening across excavations. Native micromammals provided inconsistent information about habitats and were not associated with any ritual activity or conception. The recovered reptiles, on the other hand, even though scantier than the micromammals, provided important cultural information about ritual activities of a 1st c. BC sanctuary. Such evidence is worth investigating in the future for contemporaneous materials and also for literary evidence that supports or defuses it.
18The evidence about synanthropy and imports of commensal species in the Aegean, although scanty during the 1st millennium, is incorporated in existing or new dispersal and biogeographical studies: the house mouse dispersal during Prehistory, the black rat dispersal during the Roman period and the Cretan spiny mouse import. The house mouse and the black rat, successful commensal species around the globe, are very good examples of species immigrations facilitated accidentally by humans. The spiny mouse import, an isolated insular phenomenon, shows that even small-scale events of animal circulation are worth exploring.
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Notes de bas de page
1Brothwell – Jones 1978, p. 47; Andrews 1990, p. 1. Sean Lewis is also discussing rodent dispersals and commensalism (mice and rats) based on ancient literary sources in this volume.
2Tchernov 1991, p. 156; Cucchi et al. 2020.
3Van Kolfschoten 2007, p. 2802.
4Woodburne 2006; Van Kolfschoten 2007, p. 2807.
5Geyh – Schleicher 1990, p. 315-321, 324-328, 373-377; Van Kolfschoten 2007, p. 2802.
6Doukas et al. 2018.
7Mayhew 1978.
8Stahl 1996; Brothwell – Jones 1978.
9Hofman – Rick 2018.
10Vigne 1992, p. 91, fig. 2; Vigne – Valladas 1996.
11Cucchi – Vigne 2006, p. 96.
12O’Connor 2010, p. 271.
13Hulme-Beaman et al. 2016, p. 634.
14Ibid., p. 634.
15Cucchi – Auffray – Vigne 2012.
16Tchernov 1991, p. 154-155.
17Ibid., p. 154.
18Tchernov 1984, p. 91-92.
19Shlyakov 1983.
20Hulme-Beaman et al. 2016, p. 634.
21Bonhomme – Searle 2012, p. 281-282.
22Nowadays, apart from infesting our homes, mice serve medical research; see Fox et al. 2007.
23Cucchi – Auffray – Vigne 2012, p. 66-67.
24Hamid et al. 2017.
25Cucchi et al. 2020.
26Weissbrod et al. 2017; Cucchi et al. 2020.
27Cucchi – Auffray – Vigne 2012; Cucchi et al. 2020.
28Cucchi et al. 2020.
29Ibid.; Papayiannis 2012a.
30Papayiannis 2012b, 2021.
31Honacki – Kinman – Koeppl 1982; Ervynck 2002, p. 102-103.
32Tchernov 1968, 1984.
33Ervynck 2002, p. 100-101.
34Simmons et al. 2001, p. 25, table 6.
35Armitage 1994, p. 232-233; Ervynck 2002, p. 103.
36Ervynck 2002.
37Yu et al. 2022.
38Sanges – Alcover 1980.
39Boessneck 1969.
40Audoin-Rouzeau – Vigne 1994, n. 8.
41Boessneck – von den Driesch 1975.
42Papayiannis 2012b.
43Trantalidou et al. 2019, p. 92.
44The sites are Smuszewo and Tolmicko: Teichert 1985.
45Audoin-Rouzeau – Vigne 1994, p. 132.
46Armitage 1994, p. 232-233; the absence of the black rat from the Nichoria assemblage discussed here proves this failure of establishment according to Armitage.
47Armitage – West – Steedman 1984, p. 380.
48Lortet – Gaillard 1903.
49Armitage – West – Steedman 1984, p. 380.
50Audoin-Rouzeau – Vigne 1994, p. 129, table 1.
51Sloan – Duncan 1978; Lymberakis – Mylona 2005; Lymberakis – Iliopoulos 2019.
52Mitchell-Jones et al. 1999, p. 36, 190, 262.
53Becker 1986, p. 178.
54Ibid., p. 184.
55Ruscillo 1993, p. 204, 206-207.
56Nobis 2003, p. 93-94, 98.
57Pliny, Natural history, 8.135, 30.65; Becker 1986, p. 184-185.
58Yannouli 2002, p. 321.
59Boessneck 1973, p. 25.
60Lymberakis – Mylona 2005.
61Jones 1978, p. 30.
62Sloan – Duncan 1978.
63Lymberakis – Mylona 2005.
64Lymberakis – Iliopoulos 2019.
65Payne 1995.
66Sloan – Duncan 1978.
67This species is mentioned as Microtus nivalis by Sloan – Duncan 1978, p. 75, but its taxonomic status has been updated to Chionomys nivalis and as such is mentioned in table 1.
68Mitchell-Jones et al. 1999, p. 68-69, 256-257. Two different species of shrews, C. leucodon and C. suaveolens currently inhabit the Peloponnese.
69Payne 1995, p. 289; Lymberakis – Mylona 2005, p. 299.
70Klippel – Snyder 1991, p. 180.
71Nobis 2003, p. 94, 98.
72Ruscillo 1993, p. 206-207.
73Bedwin 1992, p. 491.
74Lymberakis – Iliopoulos 2019, p. 237-238, table 5.
75Sloan – Duncan 1978, p. 75.
76Payne 1995, p. 281-284.
77Lymberakis – Mylona 2005, p. 300.
78Reese 1987, p. 258.
79Lymberakis – Iliopoulos 2019, p. 237-238, table 5.
80Reese 1989, p. 64.
81Papayiannis 2012b, 2021; Cucchi et al. 2020.
82Reumer 1986, p. 118; Ervynck 2002.
83Renaud et al. 2020, p. 610-612.
84See Cucchi 2008 for a house mouse mandible found in the Uluburun shipwreck.
85Renaud et al. 2020, p. 600, 610.
86Masseti 1998; Renaud et al. 2020, p. 610-611.
87Vigne 1999, p. 299; Horwitz – Tchernov – Hongo 2004, p. 38.
88Sloan – Duncan 1978, p. 75.
89Lymberakis – Iliopoulos 2019, p. 238.
90Ibid., table 5 and appendix.
91Mylona 2013, p. 151-153.
92Lymberakis – Iliopoulos 2019, p. 235; Mylona 2019, p. 207.
93Ibid., p. 207.
94Burkert 1993, p. 446-448; Rudloff 1999, p. 85-86, 120-123; Mylona 2019, p. 209-210.
95Aelian, On animals, 5.47; Mylona 2013, p. 158.
96Ibid., p. 157-160.
Auteur
Wiener Laboratory, American School of Classical Studies at Athens, National and Kapodistrian University of Athens
Le texte seul est utilisable sous licence Licence OpenEdition Books. Les autres éléments (illustrations, fichiers annexes importés) sont « Tous droits réservés », sauf mention contraire.
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