Keywords

1 Introduction

At present, bats (Order Chiroptera) are the second most diverse mammalian order after the rodents, with more than 1100 living species (>20% of the total extant species) distributed worldwide (apart from the poles and some isolated islands) (Simmons 2005a, b). Despite the abundant information on modern bats, their fossils are rare mostly because of their small and delicate skeleton (Gunnell and Simmons 2005). However, their distribution is universal with the oldest chiropteran fossils coming from the early Eocene of Europe (Coimbra District, Portugal), North America (Wyoming, USA), South America (Chubut, Argentina), Africa (Chambi, Tunisia or El Kohol, Algeria), Oceania (Queensland, Australia), and Asia (Gujarat, India) (Tabuce et al. 2009; Simmons et al. 2008; Tejedor et al. 2005; Sigé 1991; Ravel et al. 2011; Hand et al. 1994; Smith et al. 2007). It is evident that the appearance of Chiroptera in early Eocene is sudden, almost global, and it seems to be an isochronous event (Smith et al. 2007). Nevertheless, some of the most important questions that have yet to be answered are the temporal, geographical, and phylogenetic origins of Chiroptera.

Regarding the Greek region, recent zoological studies recorded 32 chiropteran species (Hanák et al. 2001). In the fossil record, the earliest documented chiropteran specimens come from the early Miocene locality of Lapsarna, Lesvos (Vasileiadou and Zouros 2012; Vasileiadou et al. 2017).

2 Historical Overview

The knowledge concerning the fossilized chiropteran faunas in Greece is still limited, and the Greek archive of fossilized Chiroptera is sparse. The studies so far have mainly conducted basic systematics, usually on a limited amount of specimens, and there are several factors that did not lead researchers to work with fossil Chiroptera. First, the process for the collection of microfossils requires the use of specialized equipment (sieves), unlimited water supply, etc. (for more details on the collection of microfossils and relevant information about the study of micromammals see van den Hoek Ostende 2003), which was rarely used at paleontological excavations, as researchers that worked on fossil material from the Greek region, organized excavations mainly for large mammals. Moreover, even when researchers were looking for microfossils, the chances of retrieving fossil Chiroptera were low, due to the fact that most excavations have taken place in open sites and not caves, the natural habitat for a plethora of bats. If we take all the above into account, in addition to the difficulty of the fossilization of chiropteran remains, this sparsity can be partly explained. However, the Greek archive of fossilized Chiroptera spans from early Miocene until Early/Middle Holocene. The localities where Chiroptera are documented in their faunal lists are given in Box 1.

Box 1

Early Miocene

Lapsarna, Lesvos (Vasileiadou and Zouros 2012; Vasileiadou et al. 2017)

Early/Middle Miocene

Antonios, Chalkidiki (Vasileiadou and Koufos 2005; Koufos 2006)

Late Miocene

Elaiochoria 2, Chalkidiki (Horáček 1991; Hulva et al. 2007; Sigé et al. 2014), Mytilinii, Samos (Revilliod 1922)

Miocene/Pliocene

Nea Silata, Chalkidiki (Vasileiadou et al. 2003; Koufos 2006), Maramena, Serres (Schmidt-Kittler et al. 1995; Koufos 2006)

Late Pliocene

Tourkobounia 1, Athens (Reumer and Doukas 1985)

Early Pleistocene

Tourkobounia 2, 3, 5, Athens (Reumer and Doukas 1985), Vathy, Kalymnos (Kuss 1973; Kotsakis et al. 1979)

Middle Pleistocene

Latomi 1, Chios (Storch 1975; Kotsakis et al. 1979), Petralona Cave, Chalkidiki (Sickenberg 1964, 1971; Kretzoi 1977; Kretzoi and Poulianos 1981; Horacek and Poulianos 1988; Tsoukala 1989)

Late/Middle Pleistocene

Varkiza 1, 2, Athens (van de Weerd 1973)

Late Pleistocene

Kalamakia Cave, Lakonia (Harvati et al. 2013; Darlas and Psathi 2016; Kolendrianou et al. 2020), Loutra Almopias Cave, Pella (Chatzopoulou et al. 2001; Chatzopoulou 2003, 2005; Tsoukala et al. 2006; Piskoulis 2018, 2019, 2020), Naxos, Cyclades (van der Geer et al. 2014), Kharoumes 5, Lasithi (Kuss 1970; Mayhew 1977; Dermitzakis 1977; Doukas and Papayianni 2016), Liko, Chania (Mayhew 1977; Dermitzakis 1977; Doukas and Papayianni 2016), Bate Cave, Rethymno (Kotsakis et al. 1976)

Late Pleistocene/Early Holocene

Franchthi Cave, Argolis (Stiner and Munro 2011), Anonymous Cave of Schisto at Keratsini, Piraeus (Mavridis et al. 2013)

Holocene

Charkadio Cave, Tilos (Symeonidis et al. 1973; Kotsakis et al. 1979), Vraona Cave, Attica (Symeonidis et al. 1980)

Early/Middle Holocene

Sarakenos Cave, Boeotia (Pereswiet-Soltan 2016)

Regarding the chiropteran fauna of Loutra Almopias Cave (for more information about it, see Box 2), it is worth to mention that it is currently under study within the framework of the doctorate thesis of the first author. Here are presented the first preliminary results (up to date) of the chiropteran fauna, based on fossil specimens that have been retrieved from the sediments of the cave’s floor (Fig. 1).

Fig. 1
figure 1

Finding chiropteran fossils from Loutra Almopias Cave: (a) chiropteran tooth (in circle) found in the sieved sediments, (b) screening the sieved sediments for microremains, (c) box containing cataloged chiropteran fossils, (d) over 3000 of cataloged specimens found in the sieved sediments, (e) overview of collected microremains, and (f) drawing of specimens with the use of camera lucida

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Box 2

The Loutra Almopias Cave (LAC) is located in Pella, Macedonia (Northern Greece) at the eastern slopes of Mount Voras (2524 m), about 120 km northwest of Thessaloniki. The cave is developed on the northern slope of the V-shaped gorge of Thermopotamos Stream, on the altitude of 540 m. The coordinates of the site are: N 40°58.267′, E 021°54.850′. The accumulation of sediments in the cave was in cyclic intervals (clastic and chemical sediments). The fossiliferous bed is mainly brown gravelly sand of fluvial origin. Thousands of Late Pleistocene faunal remains were collected during the excavation and through sieving procedure. The cave-site has yielded abundant faunal remains comprised of large and small mammals, birds, fish, and reptiles. The major part of the large mammalian material belongs to the cave bear (Tsoukala et al. 2006). Thirty species of rodents, insectivores, and lagomorphs have been recognized within the floor sediments providing a remarkable diversity to the LAC microfauna (Chatzopoulou 2014).

3 Phylogenetic Relationships

Chiroptera is clearly distinguished from other mammalian taxonomic groups because of their ability for powered flight and, for most of the bats, to echolocate (Gunnell and Simmons 2005; Simmons 2005b; Teeling et al. 2005). Despite the fact of being the only known mammals with the aforementioned characteristics, the monophyly of Chiroptera was largely debated since the late 1980s (Gunnell and Simmons 2005 and references therein); however, the intensification of research related to this topic and, thus, the acquirement of plethora of data (morphological, DNA hybridization and nucleotide sequence data), eventually further supported the hypothesis that all chiropteran taxa have a common (flying) ancestor (Gunnell and Simmons 2005; Teeling et al. 2005).

The aforementioned debate, resulted in the abandonment of the traditional division of Chiroptera into two sub-orders—Megachiroptera (one extant family of non-echolocating Old World fruit bats) and Microchiroptera (19 extant families of echolocating bats) (Gunnell and Simmons 2005; Simmons 2005b)—and led to the establishment of the sub-orders of Yinpterochiroptera, which includes Pteropodidae, Rhinolophidae, Hipposideridae, Megadermatidae, Craseonycteridae and Rhinopomatidae (Springer et al. 2001), and Yangochiroptera, which includes the remaining chiropteran families (Gunnell and Simmons 2005). Accordingly, echolocation in bats was evolved separately for either the two sub-orders or once with Pteropodidae (Old World fruit bats) losing it at a later stage (Teeling et al. 2005; Simmons 2005b).

As stated in the Introduction, Chiroptera appear in the fossil record during the early Eocene. New data on the phylogeny of fossil bats indicate that Onychonycteridae is the basal family for the order Chiroptera, with the rest of the extinct and extant families being sister groups (Simmons et al. 2008). The chiropteran families (20 extant and seven extinct) up to date are given in Table 1. In respect to the Greek chiropteran faunas, no phylogenetic analyses have been carried out until now. As a result, no comments can be made on the phylogeny of Greek bats.

Table 1 All chiropteran families, extinct and extant. As classification above family level is still a matter of dispute, only the most widely accepted distinct families are presented (after Gunnell and Simmons 2005; Miller-Butterworth et al. 2007; Simmons et al. 2008; Lack et al. 2010). Chiropteran families present in the fossil record of Greece are indicated with boldface
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4 Distribution

Even though the fossil record of Chiroptera is a not well-studied subject in Greece, there are quite a few localities across different regions of the country. All localities are East of Pindus Mountain Range (the absence of fossiliferous localities West of Pindus Mountain Range is because of its uplift and the absence of sedimentary basins). Out of 23 localities (Fig. 2), six are in the geographic region of Macedonia (Loutra Almopias Cave, Maramena, Antonios, Petralona Cave, Nea Silata, Elaiochoria 2), three of North Aegean Islands (Lapsarna, Latomi 1, Mytilinii), six of Central Greece (Sarakenos Cave, Anonymous Cave of Schisto at Keratsini, Tourkobounia 1, Tourkobounia 2, 3, 5, Vraona Cave, Varkiza 1, 2), two of Peloponnese (Franchthi Cave, Kalamakia Cave), one of Cyclades (Naxos) two of Dodecanese Islands (Vathy village, Charkadio Cave), and three of Crete (Liko, Bate Cave, Kharoumes 5). Further details on all localities are given in the Appendix.

Fig. 2
figure 2

Map with all the localities, where Chiroptera have been described/recorded: 1 Sarakenos Cave, 2 Vraona Cave, 3 Charkadio Cave, 4 Anonymous Cave of Schisto at Keratsini, 5 Franchthi Cave, 6 Loutra Almopias Cave, 7 Kalamakia Cave, 8 Naxos, 9 Liko Cave, 10 Kharoumes 5, 11 Bate Cave, 12 Varkiza 1 and 2, 13 Petralona Cave, 14 Latomi 1, 15 Vathy village, 16 Tourkobounia 1, 2, 3 and 5, 17 Maramena, 18 Nea Silata, 19 Mytilinii, 20 Elaiochoria 2, 21 Antonios, 22 Lapsarna. See Appendix for more information. Image exported from Google Earth Pro © 2019, map data from US Dept. of State Geographer, SIO, NOAA, U.S. Navy, NGA, GEBCO, image from Landsat/Copernicus. Scale bar equals 80 km, North faces upward

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5 Systematic Paleontology

As Chiroptera is the only known mammal group capable of self-powered flight, there are several morphological characteristics that can easily distinguish them from other extant and extinct mammals. Most importantly, their forelimbs and hind limbs are highly specialized for flight (wings) and upside-down roosting respectfully (Gunnell and Simmons 2005 and references therein). In addition to that, the chiropteran families that evolved echolocation have multiple specializations of their auditory system (Gunnell and Simmons 2005 and references therein). As described in the Introduction, complete chiropteran fossils are extremely rare, limiting their study mostly from their dental elements (including mandibles and maxillae) and humeri, which can be easily fossilized due to their harder nature (Sevilla 2016). All the above have morphological characteristics that allow paleontologists to identify fossil specimens up to species level. The identifications that follow were based on the aforementioned characteristics. Moreover, because of the limited information on most of the localities containing chiropteran fossils in Greece, this section contains information only up to the generic level. For further details on the mentioned species, see Simmons (2005a), Dietz et al. (2009), etc. Samonycteris is the only extinct chiropteran genus/species named from Greece (Revilliod 1922). It is also worthy to note that one extant chiropteran species, Myotis alcathoe , has been named from Greece (von Helversen et al. 2001).

  • Chiroptera Blumenbach, 1779Footnote 1

  • Rhinolophidae Gray, 1825

Type Genus

Rhinolophus Lacépède, 1799.

  • Rhinolophus Lacépède, 1799

Type Species

Vespertilio ferrum-equinum Schreber, 1774.

Included Taxa

Rhinolophus gr. delphinensis Gaillard, 1899, Rhinolophus ferrumequinum Schreber, 1774 (Greater Horseshoe Bat) (Fig. 3a, b), Rhinolophus ferrumequinum topali Kretzoi 1977, Rhinolophus (cf.) hipposideros Bechstein, 1800 (Lesser Horseshoe Bat) (Fig. 3c, d), Rhinolophus euryale Blasius, 1853 (Mediterranean Horseshoe Bat), Rhinolophus (cf.) mehelyi Matschie, 1901 (Mehely’s Horseshoe Bat) (Fig. 3f), Rhinolophus blasii Peters, 1866 (Blasius’s Horseshoe Bat), Rhinolophus mehelyi/euryale/blasii (Fig. 3g, h), Rhinolophus sp. (Fig. 3e), Rhinolophus sp. I (euryale group), Rhinolophus sp. II, Rhinolophus sp. III.

Fig. 3
figure 3

(a, b) Rhinolophus ferrumequinum (LAC 20600) right partial hemimandible in labial (a) and occlusal (b) view, (c, d) Rhinolophus hipposideros (LAC 16660) right partial hemimandible in labial (c) and occlusal (d) view, (e) Rhinolophus sp. (LAC 17496) right partial hemimaxilla in occlusal (e) view, (f) Rhinolophus mehelyi (LAC 16667) left partial hemimaxilla in occlusal (f) view, (g, h) Rhinolophus mehelyi/euryale/blasii (LAC 16744) left partial hemimandible in labial (g) and occlusal (h) view, (i, j) Myotis myotis (LAC 22236) left m3 in labial (i) and occlusal (j) view, (k, l) Myotis blythii (LAC 22232) left m3 in labial (k) and occlusal (l) view, (m) Myotis blythii (LAC 20468) right P4 in occlusal (m) view, (n) Myotis emarginatus (LAC 20473) left P4 in occlusal (n) view, (o) Vespertilio murinus (LAC 16379) right partial hemimaxilla in occlusal (o) view, (p) Nyctalus noctula (LAC 19325) right partial hemimaxilla in occlusal (p) view, (q, r) Barbastella barbastellus (LAC 17977) left m1 in labial (q) and occlusal (r) view, (s, t) Pipistrellus pipistrellus (LAC 18052) left C in labial (s) and occlusal (t) view, (u, v) Eptesicus serotinus (LAC 21077) right p4 in labial (u) and occlusal (v) view, (w, x) Barbastella barbastellus (LAC 16371) left partial hemimandible in labial (w) and occlusal (w) view, and (y) Miniopterus schreibersii (LAC 16383) right partial hemimaxilla in occlusal (y) view

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Comments

Rhinolophus appears in the fossil record during late Eocene–Oligocene at Quercy, France (Sigé 1978). Five Horseshoe Bat species can be found nowadays in Greece (Hanák et al. 2001; Simmons 2005a). All of those species are also present in the Greek fossil record, including R. gr. delphinensis, a fossil species that appears in the European fossil record, between MN 6 and MN 15 (Ziegler 2003). R. gr. delphinensis is present in Elaiochoria 2, R. ferrumequinum and R. hipposideros are present in Sarakenos Cave, R. hipposideros and R. blasii are present in Charkadio Cave, R. ferrumequinum, R. hipposideros, R. euryale, R. mehelyi, R. blasii, R. mehelyi/euryale/blasii, and Rhinolophus sp. are present in Loutra Almopias Cave, Rhinolophus sp. is present in Kalamakia Cave, R. ferrumequinum topali, R. cf. hipposideros, R. cf. mehelyi, Rhinolophus sp. I (euryale group), II and III are present in Petralona Cave.

  • Vespertilionidae Gray, 1821

Type Genus

Myotis Kaup, 1829.

Comments

Vesper Bats appear in the fossil record during early Eocene (Miller-Butterworth et al. 2007). Several species have been identified in the Greek fossil record at different taxonomic levels. Vespertilionidae sp.1 and sp.2 are present at Nea Silata.

  • ★Samonycteris Revilliod, 1922

Type Species

★Samonycteris majori Revilliod, 1922.

Included Taxa

Samonycteris majori Revilliod, 1922.

Comments

Major (1891) assigned an incomplete chiropteran skull retrieved from the late Miocene Mytilinii basin (Samos) as Chiroptera indet. Later on, Revilliod (1919) refers to this specimen for the first time as S. majori, while its description was published in 1922 by the same author. Samonycteris has a single robust upper incisor and its skull has a morphological resemblance to that of Otonycteris and/or Eptesicus, which, however, differs in several morphological features (e.g., the cochlea) and is also a little smaller than the latter (Revilliod 1922; Horáček 1991). S. majori is a monotypic taxon.

  • Myotis Kaup, 1829

Type Species

Vespertilio myotis Borkhausen, 1797.

Included Taxa

Myotis myotis Borkhausen, 1797 (Greater Mouse-eared Bat) (Fig. 3i, j), Myotis blythii Tomes, 1857 (Lesser Mouse-eared Bat) (Fig. 3k–m), Myotis blythii oxygnathus Monticelli, 1885, Myotis bechsteinii Kuhl, 1817 (Bechstein’s Bat), Myotis emarginatus Geoffroy, 1806 (Geoffroy’s Bat) (Fig. 3n), Myotis capaccinii Bonaparte, 1837 (Long-fingered Bat), Myotis cf. daubentonii Kuhl, 1817 (Daubenton’s Bat), Myotis mystacinus? Kuhl, 1817 (Whiskered Bat), Myotis myotis/blythii, Myotis sp., Myotis sp. I, Myotis sp. II, cf. Myotis (small sized).

Comments

Myotis appears in the fossil record during early Oligocene (Gunnell et al. 2017). Ten to eleven Mouse-eared Bat species currently inhabit the Greek region (Hanák et al. 2001), and some of them are also present in the fossil record. M. blythii and M. myotis/blythii are present in Sarakenos Cave, Myotis sp. is present in Vraona Cave, M. blythii and Myotis sp. are present in Charkadio Cave, M. cf. blythii is present in Kalamakia Cave, M. myotis, M. blythii, M. bechsteinii, M. emarginatus, M. capaccinii, M. mystacinus?, M. myotis/blythii and Myotis sp. are present in Loutra Almopias Cave, M. myotis, M. blythii ssp., M. blythii oxygnathus, M. emarginatus, M. cf. daubentonii, Myotis sp. I and II are present in Petralona Cave,Footnote 2 cf. Myotis (small sized) is present in Elaiochoria 2.

  • Nyctalus Bowdich, 1825

Type Species

Nyctalus verrucosus Bowdich, 1825 (= Vespertilio leisleri Kuhl, 1817).

Included Taxa

Nyctalus lasiopterus? Schreber, 1780 (Greater Noctule Bat), Nyctalus leisleri Kuhl, 1817 (Leisler’s Bat), Nyctalus noctula Schreber, 1774 (Noctule Bat) (Fig. 3p).

Comments

Nyctalus appears in the fossil record during the Oligocene (Horáček 2001). Three Noctule Bat species can be found nowadays in the Greek region (Hanák et al. 2001; Simmons 2005a). In the fossil record of Greece, N. noctula is present in Sarakenos and Loutra Almopias Caves, N. cf. noctula is present in Petralona Cave, N. lasiopterus? and N. leisleri are present only in Loutra Almopias Cave.

  • Pipistrellus Kaup, 1829

Type Species

Vespertilio pipistrellus Schreber, 1774.

Included Taxa

Pipistrellus pipistrellus Schreber, 1774 (Common Pipistrelle Bat) (Fig. 3s, t), Pipistrellus(?) sp.

Comments

Pipistrellus appears in the fossil record during Middle Pleistocene (Horáček and Jahelková 2005). Four Pipistrelle Bat species can be found nowadays in the Greek region (Hanák et al. 2001; Simmons 2005a). In the Greek fossil record, P. pipistrellus is present in Loutra Almopias Cave, Pipistrellus(?) sp. is present in Petralona Cave, and Pipistrellus sp. is present in Kalamakia Cave.

  • Hypsugo Kolenati, 1856

Type Species

Vespertilio savii Bonaparte, 1837.

Included Taxa

Hypsugo savii Bonaparte, 1837 (Savi’s Pipistrelle Bat).

Comments

Hypsugo was included in Pipistrellus until mid-1980s, however, since then it gained separate generic status (Horáček and Hanák 1985). One Asian Pipistrelle Bat species can be found nowadays in the Greek region (Hanák et al. 2001), which is also present in the Greek fossil record. H. savii is present only in Petralona Cave.

  • Vespertilio Linnaeus, 1758

Type Species

Vespertilio murinus Linnaeus, 1758.

Included Taxa

Vespertilio murinus Linnaeus, 1758 (Parti-colored Bat) (Fig. 3o).

Comments

Vespertilio appears in the fossil record during the Pliocene (Gunnell and Simmons 2005). One Parti-colored Bat species can be found nowadays in the Greek region (Hanák et al. 2001), which is also present in the Greek fossil record. V. murinus is present in Loutra Almopias and Petralona Caves.

  • Eptesicus Rafinesque, 1820

Type Species

Eptesicus melanops Rafinesque, 1820 (= Vespertilio fuscus Beauvois, 1796).

Included Taxa

Eptesicus serotinus Schreber, 1774 (Serotine Bat) (Fig. 3u, v), Eptesicus sp.

Comments

Eptesicus appears in the fossil record during the early Miocene (Gunnell and Simmons 2005). Three Serotine Bat species can be found nowadays in the Greek region, with E. bottae only present in Rhodes (Hanák et al. 2001; Simmons 2005a), and at least one of them is also present in the Greek fossil record. E. serotinus is present in Loutra Almopias and Sarakenos Caves, Eptesicus sp. is present in Petralona Cave.

  • Barbastella Gray, 1821

Type Species

Vespertilio barbastellus Schreber, 1774.

Included Taxa

Barbastella barbastellus Schreber, 1774 (Western Barbastelle Bat) (Fig. 3q, r, w, x).

Comments

Barbastella appears in the fossil record during the Early Pleistocene (Rydell and Bogdanowicz 1997). One Barbastelle Bat species can be found nowadays in the Greek region (Hanák et al. 2001), which is also present in the Greek fossil record. B. barbastellus is present only in Loutra Almopias Cave.

  • Plecotus Geoffroy, 1818

Type Species

Vespertilio auritus Linnaeus, 1758.

Included Taxa

Plecotus austriacus Fischer, 1829 (Gray Long-eared Bat), P. auritus/austriacus.

Comments

Plecotus appears in the fossil record during the middle Miocene (Gunnell and Simmons 2005). Three Long-eared Bat species are currently present in the Greek region (Hanák et al. 2001). P. austriacus is present in the fossil record of Charkadio Cave and P. auritus/austriacus is present in the fossil record of Loutra Almopias Cave.

Type Genus

Miniopterus Bonaparte, 1837.

  • Miniopterus Bonaparte, 1837

Type Species

Vespertilio ursinii Bonaparte, 1837 (= Vespertilio schreibersii Kuhl, 1817).

Included Taxa

Miniopterus schreibersii Kuhl, 1817 (Schreibers’ Bent-winged Bat) (Fig. 3y).

Comments

Miniopterus appears in the fossil record during middle Miocene (Horáček et al. 2013). One Bent-winged Bat species can be found nowadays in Greece (Hanák et al. 2001), which is also present in the Greek fossil record. M. schreibersii is present in Loutra Almopias Cave, Petralona Cave, and Vathy village.

  • Rhinopomatidae Bonaparte, 1838

Type Genus

Rhinopoma Geoffroy, 1818.

  • Rhinopoma Geoffroy, 1818

Type Species

Vespertilio microphyllus Brünnich, 1782.

Included Taxa

Rhinopoma aff. hardwickii Gray, 1831 (Lesser Mouse-tailed Bat).

Comments

The Mouse-tailed Bats (R. aff. hardwickii) appear in the fossil record during Late Miocene at Elaiochoria 2, Greece, which is the earliest and sole occurrence of this family in Europe (Hulva et al. 2007).

  • Megadermatidae Allen, 1864

Type Genus

Megaderma Geoffroy, 1810.

Included Taxa

cf. Megadermatidae gen. and sp. indet. (False Vampire Bats).

Comments

The False Vampire Bats appear in the fossil record during late Eocene (Agnarsson et al. 2011). Fossils of Megadermatidae have been retrieved from several European sites (Gunnell and Simmons 2005), including Greece. cf. Megadermatidae gen. and sp. indet. is present in Antonios.

  • Chiroptera indet.

Comments

Undetermined species of Chiroptera (also mentioned as “Chiroptera,” “Bats,” etc.) have been found at Anonymous Cave of Schisto at Keratsini, Franchthi Cave, Loutra Almopias Cave, Naxos, Liko, Kharoumes 5, Bate Cave, Varkiza 1, 2, Latomi 1, Tourkobounia 1, 2, 3, 5, Vathy village, Maramena, and Lapsarna.

6 Concluding Remarks

It is evident that the fossil record of Chiroptera is not well-studied in Greece, with sparse finds and most of the works reaching up to generic level. Nevertheless, the following taxa have been identified in Greece: Rhinolophus gr. delphinensis, Rhinolophus ferrumequinum, Rhinolophus ferrumequinum topali, Rhinolophus (cf.) hipposideros, Rhinolophus euryale, Rhinolophus (cf.) mehelyi, Rhinolophus blasii, Rhinolophus mehelyi/euryale/blasii, Rhinolophus sp., Rhinolophus sp. I (euryale group), Rhinolophus sp. II, Rhinolophus sp. III, Samonycteris majori, Myotis myotis, Myotis (cf.) blythii (ssp.), Myotis blythii oxygnathus, Myotis bechsteinii, Myotis emarginatus, Myotis capaccinii, Myotis cf. daubentonii, Myotis mystacinus?, Myotis myotis/blythii, Myotis sp., Myotis sp. I, Myotis sp. II, cf. Myotis (small sized), Nyctalus lasiopterus?, Nyctalus leisleri, Nyctalus (cf.) noctula, Pipistrellus pipistrellus, Pipistrellus(?) sp., Hypsugo savii, Vespertilio murinus, Eptesicus serotinus, Eptesicus sp., Barbastella barbastellus, Plecotus austriacus, Plecotus auritus/austriacus, Miniopterus schreibersii, Rhinopoma aff. hardwickii, cf. Megadermatidae gen. and sp. indet., Vespertilionidae sp.1, Vespertilionidae sp.2, and Chiroptera indet.

Rhinopoma aff. hardwickii from Late Miocene site Elaichoria 2 is the earliest and sole occurrence of Rhinopomatidae in Europe and consequently Greece. Furthermore, the study of Chiroptera from Loutra Almopias Cave, which is presented here for the first time as an updated preliminary list of taxa, will give us the first complete chiropteran record in the Greek region. In addition, the revision of the existing specimens retrieved from the localities mentioned herein might allow us to reach to a more detailed identification. In any case, the acquisition of new material and basic systematic study will give us the opportunity to carry out more targeted research on this relatively unexplored topic (i.e., phylogenetic analysis) that will allow us to extent our knowledge on the chiropteran evolution in the Balkan Peninsula.