Middle Pleistocene bats (Mammalia: Chiroptera) from the Yarimburgaz Cave in Turkish Thrace (Turkey)

Th e Yarimburgaz Cave near Bosporus is an important Middle Pleistocene fossil site both in terms of Archeology and Paleontology in the northeast Mediterranean, containing the richest bat fauna of that time. Th ere are at least 14 and possibly as many as 16 species of the genera Rhinolophus, Myotis, Plecotus, and Miniopterus present in this locality. Th e identifi cation is based on the morphology of skulls, jaws, and humeri. Th e cave contains three sedimentary cycles. Cycle I is the oldest unit and produces most of the species, eight, with Rhinolophus mehelyi and Myotis blythii the most frequent. Th e presence of these species suggests a Mediterranean climate with cool, rainy winters and hot, moderately dry summers (K  V 2001). In cycle II very few fossils were found. Cycle III contains fewer individuals than cycle I with only six taxa represented. Myotis blythii and Miniopterus schreibersi are the dominating species. Given the presence of the rodents and lagomorphs of the Yarimburgaz Cave, a heterogeneous climate with changing colder, dryer, but also warmer phases which are indicated by the bats, can be supposed. [Mittelpleistozäne Chiroptera (Mammalia) aus der Yarimburgaz-Höhle in Türkisch Th razien (Türkei)] Kurzfassung: Die Yarimburgaz-Höhle am Bosporus ist die wichtigste Fundstelle für die mittelpleistozäne Archäologie und Paläontologie im nordöstlichen Mittelmeerraum und beinhaltet in diesem Zeitraum die reichste Fledermausfauna der Region. Aus der Lokalität werden hier die Chiropteren anhand von Crania, Mandibulae und Humeri untersucht. Es sind mindestens 14, eventuell bis zu 16 Arten aus den Gattungen Rhinolophus, Myotis, Miniopterus und Plecotus vorhanden, wodurch die bisherige Fossilfauna um weitere neun bis elf Arten ergänzt wird. Von den drei Sedimentationszyklen enthält der älteste, Zyklus I, die meisten Fledermausfunde. Unter den acht verschiedenen Arten sind Rhinolophus mehelyi und Myotis blythii am häufi gsten vertreten und weisen auf mediterranes Klima mit kühlen, regnerischen Wintern und heißen, trockenen Sommern hin (K  V 2001). Aus Sedimentationszyklus II sind außer einer Mandibel von R. mehelyi keine weiteren Funde überliefert. Zyklus III beinhaltet mit insgesamt sechs Taxa wesentlich weniger Funde als Zyklus I. Darunter sind Myotis blythii und Miniopterus schreibersi die dominierenden Arten. Verglichen mit den Rodentia und Lagomorpha aus der Yarimburgaz-Höhle, die als Einwanderer des südrussischen Steppengürtels mehrere kälteangepaßte Arten umfassen und rezent nicht mehr in Th razien vorkommen, kann ein inhomogenes Klima angenommen werden, mit wechselweise kühleren, trockeneren, aber auch wärmeren Phasen, welche durch die Fledermäuse angedeutet werden.

Hannover 2005 dle Pleistocene, predominantly from the upper two meters of sediments from the entrance area of the lower cave (H  A ; K ).Large mammals are also excavated with the cave bear Ursus deningeri being the most frequent.Within the carnivores Vulpes, Canis, Panthera, Felis, and Crocuta are also present.Th e herbivores contain Equus, Sus, Dama, Cervus, Megaloceros, Capreolus, Bos/ Bison, Gazella, and Capra (S ).Th is fauna may represent the hunting activity of Early man, but others species like the cave bear most probably used the cave for hibernation.Th e small mammal fauna is not as rich and represented by some Insectivora, Rodentia (i.a.Scioridae, Murinae, Cricetinae, Arvicolinae), and Lagomorpha (S  K ).Bats make up the biggest portion within the small mammals.Ecological conclusions will be drawn on the basis of recent exponents ecology.
Here we aim to interpret the paleoclimatic conditions present and these will be compared with the ecological data derived from the Rodentia and Lagomorpha S  K () described from the same locality.Th is will allow us to test if both faunal groups indicate the same climatic conditions.As bats are rare in the fossil record this Middle Pleistocene is a highly signifi cant location.

Locality
Th e Yarimburgaz Cave is located 20 km west of Istanbul in Turkish Th race and belongs to a vast complex developed in Middle Eocene of Lutetian limestone.Th e cave system contains two chambers, the lower one continues for about 500 m the other for some 50 m.Th ese two chambers are connected through a ramp like passage but each has its own entrance (fi g. 1).For a long time part of the cave was used for various purposes.First scientifi c explorations were established during [1964][1965].In 1986 M. Ö made a systematic survey in the upper cave.During 1988-1990 C. H andG.A carried out the main excavations in the lower chamber, producing the bat fauna described here.All excavations were pursued in the entrance area.Th e sediments fi lling the lower cave were excavated in various unconnected trenches (P-90, S-89, T-89, A-71 & V-88, Y-88, U-88, R-90, and Z-88) roughly following the midline of the cave from north to south (fi g. 1; 2).Th ese trenches were designated with a capital letter followed by the number of the year the excavation took place, except for trench A-71, which was excavated by Ö in 1986.Th e letters Q, W, and X do not occur because they do not exist in Turkish (F  MM 1997).
Th e level numbers in each trench were assigned by the excavators independently of those in the other trenches, except for trenches S-89 and T-89, which were dug simultaneously (F & MM 1997).Th erefore a correlation of the diff erent levels was not possible and there are no data about the thickness of the levels.However, the excavation levels were grouped by F & MM (1997) into lithostratigraphic units designated Stratum R through Stratum Z (from bottom to top) from three sedimentary cycles (fi g. 2, tab.1).In this case, additional letters W and Z were used, so one has to be certain the letter used for classifi cation are correctly assigned to either the excavation square or to a stratigraphic unit.Only square U could not be correlated because it was very shallow, over bedrock and its stratigraphy was not informative (F & MM 1997).Th e deepest trench, A-71 & V-88, was dug fi ve meters below the surface without reaching bedrock.However below two meters no artifacts or bones were found (F  MM 1997).
Depending on the position of the square there variations can be found in the composition of Middle Pleistocene bats from the Yarimburgaz Cave in Turkish Th race (Turkey)  G 1998).Generally, the fi rst three are called summer roosts and can be in tree hollows or caves, whereas hibernating bats are normally found in caves.Some of the chiropterans may have used the cave seasonally, only in summer, whereas others may have used the cave to For climatic reconstruction it would be useful to separate species occurring only during summer from those present year round.However, such a discrimination cannot be done in the fossil record.
Recent Microchiroptera are distributed world wide with 17 families, 150 genera, and 808 species, and are absent only in polar regions and on some isolated oceanic islands (N 1999).
Most are insectivorous, but in the tropics some are food specialists, which feed on fruits, leaves, nectar, pollen, fi shes, other small vertebrates, or blood (K , N ).Both infraorders of microchiropterans from Yarimburgaz Cave could be verifi ed, Yinochi-roptera (premaxillae almost never fused with maxillary) and Yangochiroptera (premaxillaries always fused with maxillaries in adults).Th e fi rst is represented by the family Rhinolophidae, including four species of one genus (tab.2).Th e Yangochiroptera are represented by three genera and ten to twelve species of the Vespertilionidae (tab.2).Th e material consists of 229 fragmentary rostra and maxillae, 1562 mandibles, and 717 humeri were identifi ed.Fig. 3 provides a general impression of the preserved material.Light sedimentary incrustation of the studied objects was removed with an insect needle.Further material is damaged and identifi cation beyond Microchiroptera is obviated.Apart from the humeri no further postcranial material was analyzed,    (1959) in this paper the eutherian dental formula is modifi ed to I 1 (2)/i 1 2 (3), C 1/c 1, P 1 (3) 4/p 1 (3) 4, M 1 2 3/m 1 2 3. Th e teeth of the positions in brackets can be absent.

Determination of the material
Rhinolophidae G,  Horseshoe Bats Rhinolophus L  Th ere is a single genus -Rhinolophus -with 64 living species (K ).Th e nasal region of the skull is clearly bulged out.Th e premaxilla builds a fragile intermaxilla which is only linked to the palatinum by connective tissue.Th erefore the intermaxilla cannot be conserved in fossil skulls but remains a huge palatine incision.Th e mandible is conspicuously long and fl at, the most distinguishing character is the extremely fl at processus (proc.)coronoideus.Th e fi rst teeth within the upper jaw are the large, strongly curved canines as the incisivi situated on the palatine are not preserved.Th e P1 is mostly reduced and placed buccally.Th e P4 is large and distally has a sharp crista.
C L On M1-2 the hypocone is well developed and curved in mesio-distal direction.Th e M 3 is slightly smaller than the M2, the reduction aff ects mainly metastyle and hypocone.In the lower jaw the incisivi are closely spaced.Th eir crown is three-parted and spatulate.Th e c is very high and surrounded by a broad cingulum.Two lateral ridges frame a broad concave and distal directed plane.Th e p1 is relatively small.Th e p3 is maintained only as a diminutive tooth at the buccal outside of the tooth row.In opposite, the p4 is relatively large and builds three distinct ridges.Th e morphology of m1-m3 is nyctalodont (fi g. 4).Th e distal epiphysis of the humerus is characterized by a long spikelike processus styloides as well as by a very broad expanded epitrochlea and epicondylus.A deep incisur is located between trochlea and condylus.Rhinolophus ferrumequinum (S ), Greater Horseshoe Bat cycle I: 1 rostrum, 4 mandibles, 11 humeri; cycle III: 2 maxillae, 1 mandible, 1 humerus; uncorrelated: 1 rostrum, 3 maxillae, 14 mandibles, 5 humeri R. ferrumequinum is the largest European Rhinolophus-species and it can easily be identifi ed by its dimensions.. Th e P1 of the upper jaw is very small and shifted buccally so the canine and the P4 are in contact.Similar to that the p3 in the lower jaw is enormously reduced and displayed buccally beyond the tooth row.Th e molars are nyctalodont (fi g. 4).Th e distal epiphysis of the humerus has a width  Abb.6: Untere Zahnreihe von Rhinolophus euryale und Rhinolophus blasii.Bei R. euryale nimmt der p1 weniger als 50% der Fläche des p4 ein, bei R. blasii liegt der Anteil der Fläche über 50% (nach S  G 1998).
C L of about 5 mm between epitrochlea and epicondylus which is found in no other rhinolophid species.
Rhinolophus euryale B , Mediterranean Horseshoe Bat cycle I: 4 humeri; uncorrelated: 1 humerus Th e most striking characteristic on the skull is an ossifi ed bar between the foramen (for.)infraorbitale and the orbita.Th is bar is short and relatively broad (fi g. 5).Th e P1 is located buccally at the border of the tooth row and is constricted by the spiky C and the P4.Th e M3 is slightly reduced.In the lower jaw the crownplane of the p1 achieves a maximum of 50% of the plane of the p4 (fi g. 6).Th e p3, which lies between them, is shifted buccally far beyond the tooth row.Its peak reaches the cinguli of the remaining premolars.
On the distal epiphysis of the humerus an incisur is situated at the transition of the proc.styloides to the epitrochlea in cranial and caudal view (fi g. 7).Th e proc.styloides is relative slender.In the same views the trochlea forms a distinct edge towards distal.Looking at the caudal outline of the trochlea no sharp edge is developed in lateral view.
Because there are great analogies of R. euryale with and R. mehelyi concerning characteristics and size only fi ve humeri could be clearly identifi ed.Th is problem will be discussed after the description of R. blasii and R. mehelyi and similar species will be combined as one group.
Rhinolophus blasii P , Blasius's Horseshoe Bat cycle I: 1 mandible, 8 humeri; uncorrelated: 3 humeri Th e skull exhibits a smooth occipital depression.Th e ossifi ed bar between the for.infraorbitale and the orbita is relatively short and narrows slightly to the middle.Th e P1 and P4 are often separated through a diastema.In the lower jaw the crown-plane of the p1 takes not less than 50% of the plane of p4 (fi g. 6).Th e p3 in-between is squashed in and strongly reduced.Th e top of the proc.styloides on the distal epiphysis of the humerus is rounded towards medial (fi g. 8).Th e transition of the proc.styloides towards the epitrochlea is shallow in cranial and caudal view.Based on the smaller width of the epiphysis at an average of 4 mm in addition to the other characteristics the humeri of R. blasii can easily be identifi ed.
Only one lower mandible could be recognized for R. blasii.Here the p4 is still preserved and   Many species from the temperate areas migrate between summer-and winter roosts depending.
In the Yarimburgaz Cave this could be demonstrated for the genera Myotis, Plecotus, and Miniopterus.
Usually two incisors are situated on both sides of the intermaxilla of the skull of vespertilionids.Th e incisura nasalis lies deep in the premaxilla and reaches to the canine, maximum to the fi rst premolar.Th ere is no bulb of the nasal region as in rhinolophids.Th e lower jaw is characterized by a high and compact proc.coronoideus (fi g. 12).Myotis K 1829, Little Brown Bats Within the chiropterans Myotis is the most widespread form being not found only in the arctic, subarctic and antarctic regions as well as on many oceanic islands (N, 1994).With 84 recent species this genus are also the most diverse within the bats (K, ).
Th e rostrum of the relatively slender skull is elongated.Th e inc.nasalis is rounded dorsally.Some species build a crista sagittalis but the linea postorbitalis is never prominent.Th e for.infraorbitale is situated above the P4/M1 and is separated from the orbita by a broad ossifi ed bar.Th e cr.masseterica and the fossa masseterica distinctly emerge.Th e foramen mentale is situated under c/p1, rarely under p1.Th e upper incisivi build two to three peaks of nearly the same height.Th ere is a diastema between the I2 and the caninus.Th e latter is relatively small but massive and with a sharp distal crista surrounded by a broad cingulum.Th e P1 and P2 are tiny, whereas the following P4 nearly reaches the height of the C and surmounts the peaks of the molars.Th e molars are dilambdodont and lack any peculiar characteristics.Th e lower incisors are closely spaced with small overlap.Th ey are spatulate; their crowns develop three to four peaks.Th e i3 is signifi cantly larger than i1 or i2 and forms a pronounced inner protuberance.Th e c is slightly curved backwards.A bucco-distal and a lingual ridge build a concave distal plane.Th e p4 is the tall-  Myotis bechsteini (K 1817), Bechstein's Bat cycle I: 33 mandibles; cycle II: 2 humeri; cycle III: 1 maxilla, 2 mandibles; uncorrelated: 1 rostrum, 13 maxillae, 94 mandibles, 1 humerus Th e P1 of the upper jaw is as large as or larger than the P3, and the latter is situated within the tooth row.Th ere is often a diastema between P3 and P4.Additional cones, called protocoluli, can be found at the mesial inner plane of M1-3.If they are present, they are only weakly developed.Th e proc.coronoideus of the lower jaw is very high, its cr.masseterica nearly arises at right angle (fi g. 12).Th e distal epiphysis of the humerus can easily be identifi ed by the missing proc.styloides (fi g. 14).In cranial view the rostral end of the trochlea does not reach the rostral end of the epitrochlea proximally.Th e width of the epiphysis of 2,9-3,1 mm is a further characteristic of the relatively large species (F al. 1973).In fi g. 15 the lower tooth row of the Bechstein's Bat from Yarimburgaz is compared with recent and Early Pleistocene material from Deutsch-Altenburg (DA 28)of this species.As the fossil record of DA 28 fi ts in the variability of Yarimburgaz there is no variation of size since the Early Pleistocene.Th e recent values do not cover the amplitude of variation.
Myotis nattereri (K 1817), Natterer's Bat uncorrelated: 2 maxillae, 2 mandibles Th e arcus zygomaticus of the skull is slender (fi g. 16a).In lateral view the fi rst two upper incisive diverge, causeing a relatively large angle between both tooth crowns.Th e C is comparatively small.Th e P1 is insignifi cantly larger than the following P3; both are highly reduced.
Regarding the upper teeth, the c of the lower jaw is relatively small and the proportion of p1 to p3 is concordant with that of the upper jaw (fi g. 16a).Th e cross section of the c is oval.
As there are no specifi c characteristics of the middle-sized humeral epiphysis (2,6-2,7 mm) (F et al. 1973).M. emarginatus and M. daubentoni are of equal size and thus no humeri of these dimensions could be referred to a specifi c species.Th ere is a lot of overlap in morphological and biometrical characteristics of M. nattereri with M. emarginatus which will be discussed later.Myotis emarginiatus, (G 1806), Geoffroy's Bat cycle I: 1 rostrum, 1 maxilla; uncorrelated: 5 mandibles Th e arctus zygomaticus is clearly broadened in lateral view (fi g. 16b).Th e crowns of the incisivi are parallel.Th e C is long and acuminate.Th e P1 and P3 are reduced, the P3 reaches maximum half of P1 (fi g. 16b).M1-3 develop no protoconuli.Th e i2 of the lower jaw develops four peaks and a further inner conulus can be  C L present.Th e cross section of the large c is round.Th e p1 is slightly larger than the p3 (fi g. 16b).
As mentioned before there are no characteristics which can be attributed to the humerus.Th e C is rather compact.Th e P4 is signifi cantly larger than the tiny P1 and P3 and it reaches across the peaks of the molars.In the mandi-ble the i1 and i2 are three-coned; i3 developed four peaks and is substantially widened.Th e c is only weakly developed.As in the upper jaw the crown of the p4 overtops the peaks of the other teeth.
In lateral view the proc.styloides protrudes caudally above the shank of the humerus (fi g. 17).
A distinct groove, which curves from caudal to lateral, separates the proc.styloides from the trochlea.Another distinguishing feature is the small size of the humerus with an epiphyseal width of 2,7 mm (F et al. 1973).Th e material referred to M. capaccinii also comprises a very small humerus that shows the characters described above but has an epiphyseal diameter of only 2.38 mm.Th is humerus most likely belongs to a juvenile animal.Plecotus G, 1818, Old World Longeared Bats Th is genus is distributed in Europe, Asia, North-Africa and America (Kanada to Mexico) with eight recent species (K, 1994; N, 1994).Th e profi le of the skull is relatively fl at; the frontal region is mildly bulged.A distinct cr.lacrimalis is situated on the rostral edge of the orbita.Th e bullae tympanic are large; the distance between them is considerably smaller than their individual diameter.With the reduction of one premolar the dental formula is 2/3, 1/1, 2/3, 3/3.Th e fi rst of the two upper incisivi is considerably larger and buccally has a further conus.Usually a diastema is situated between the highly reduced P1 and the P4.Th e cingulum of the latter can build a mesio-lingual protoconus.Th e molars show no diagnostic features except for the greatly reduced M3.Within the mandibular dentition the inicivi have three to four peaks and the i3 is slightly widened.Th e pronounced cingulum of the c is mesial extended and builds an additional protuberance on half level of the tooth height.Th e p3 is narrowed by the p1 and p4.Th e morphology of the molars is myotodont (fi g 4).

C L
In cranial view the transition of the trochlea to the condylus at the distal epiphysis of the humerus is clearly convex at the articulated joint.In the same view the rostral edge of the trochlea does not reach the rostral edge of the epitrochlea.Trochlea and condylus overtop the proximal peak of the epicondylus in lateral view.Caudally the fossa olecrani is not present or only weakly developed.
Plecotus auritus (L 1758), Brown Bigeared Bat uncorrelated: 1 maxilla, 1 mandible Th e most prominent feature of the skull is the arcus zygmaticus, which is highly bulged in its middle.Th e C is rather short.Th e lower jaw can be identifi ed by the thickened and rounded end of the proc.angularis (fi g. 18a).Similar to the upper jaw the c is relatively short.At the distal epiphysis of the humerus the transition of the epitrochlea to the proc.styloides is angular (fi g. 19a).Th e average width of the epiphysis is about 2.7-2.9 mm (F et al. 1973).
Plecotus austriacus (F 1829), Gray Bigeared Bat uncorrelated: 4 mandibles Th e arcus zygmaticus on the skull is slightly broadened in its middle but it becomes signifi cantly slender towards aboral.Th e C is more slender and more pointed than in P. auritus.Th e proc.angularis at the lower jaw ends blunt with a distinct projection at its anterior third (fi g. 18b).Th e c shows a shallow bending towards distal.
Th e transition of the epitrochlea to the proc.styloides at the distal epiphysis of the humerus is not as angular as in P. auritus in caudal view (fi g. 19b).Th e average width of the epiphysis is also about 2.7-2.9 mm (F et al. 1973).
group Plecotus auritus-austriacus cycle III: 1 maxilla, 1 mandible, 1 humerus; uncorrelated: 1 maxilla, 29 mandibles, 4 humeri Th e diff erences these two species are very marginal.Th e diagnostic characters described above are mainly based on structures which are prone to damage during the fossilization process.Other characters, in particular the distal epiphysis of the humerus, are variable and cannot be used on its own (fi g. 19).Th e shape of the proc.angular of P. auritus often appears in P. austriacus as personal observation of comparison material revealed.Th erefore, it is not possible to precisely separate the species.Miniopterus B 1837, Long-winged Bats Th irteen species of this genus are distributed world-wide except for America and the arctic Th e humerus can be determined with some certainty by the characteristic shape of the distal epiphysis.Additional to these the epiphysis is relatively narrow with an average width of 2.7 mm (F et al. 1973).

Composition of the bat fauna
Th ree sedimentary cycles were diff erentiated by H  A (1989) and F  MM (1997).A rich bat fauna originates from cycles I and III and each of these units is summarized here.Th e denoted number of the taxa is the minimal number of individuals (MNI) of lower jaws identifi ed.For Rhinolophus euryale the humeri were counted because there were no mandibles (fi g. 21).Due to the fact that there is a lot of overlap in the characteristics of Myotis mystacinus, M. brandti, and M.  In addition, square U off ered a great number of fossil bat remains which cannot be precisely correlated to these cycles and are handled separately here.Th e genus Myotis is represented by seven to nine species.However, the most frequent species is Rhinolophus mehelyi which represents 52% of the total sample of bats (fi g. 21).

Stratigraphic position of the three cycles and trench U
No exact stratigraphic position can be given for the three cycles.Th e excavation levels were designated into lithostratigraphic units based on diff erent layers of erosion (fi g. 2).(S 1994).Th e signifi cant steppic infl uence is indicated by immigrants from the Ukraine which are no longer found in the area today: Cricetus cricetus, Cricetulus migratoris, Lagurus transiens, Sicista subtilis, and Ochotona pusilla.Although the rodents characterize a harsher climate than the bats, there is no indication that the bats come from other layers than the rodents.Most likely the chiropterans as well as rodents and lagomorphs coexisted in the area.Comparing the recent distribution of these species with that of the chiropterans, all taxa share parts of their ranges.Only the two thermophilic species R. mehelyi and M. schreibersi do not fi t in the same ecological surrounding nor overlap in geographic range with Cricetus cricetus, Lagurus transiens, and Ochotona pusilla (E; L- & F 1985).Th erefore, one has to assume that these bats used the cave seasonally and came here in summer only or used it for hibernation.S (1994) postulated an open landscape for cycle III with a signifi cant steppic infl uence.Th e bat fauna represents an open landscape too, but not as cold and dry.Th e time involved in cycle III might represent a heterogeneous climate with changing colder, dryer, and warmer phases which is not fully deducible from the coarse stratigraphic allocation of the fossil material.Due to changes in the climate the history of the bat distribution might be characterized by a local extinction and reinvasion of taxa which cannot be determined from the sediments of the Yarimburgaz Cave.

Trench U
Th e sediments of the isolated square U could not be correlated with one of the three cycles because it was very shallow over bedrock and its stratigraphy not informative as mentioned before (H  A ; F  MM ).Almost 70% of the bats originate from this square.Interestingly the majority of the other small mammals (78%) was also concentrated within square U (S C L 1994).Looking at the map (fi g. 1) this huge concentration within a single square seems not to refl ect the biotope.As Yarimburgaz Cave is still an active karst-system it may be possible that the small mammals were concentrated by transportation mixed up by various sources.Th e uninformative stratigraphy made a correlation to any layer of the other squares impossible.
Presumably the accumulation of bats represents diff erent time periods which could not be separated during excavation.Th erefore, these very rich layers cannot be contributed to the reconstruction of the climatic history of the Yarimburgaz Cave.13-15 species of bats were identifi ed:

Fig. 1 :Fig. 2 :
Fig. 1: Map of the entrance area of Yarimburgaz Cave with openings to the upper and lower chamber including the squares (after F  MM ).Abb.1: Karte des Eingangsbereiches der Yarimburgaz-Höhle mit den Öff nungen zur oberen und unteren Kammer und Grabungsquadraten (nach F  MM ).

Tab. 2 :
Systematic classifi cation of the identifi ed chiropterans based on MK  B ().Tab.2: Systematische Gliederung der untersuchten Chiropteren nach MK  B (2000).Tab.1: Correlation of the excavation levels grouped into lithostratigraphic units designated Stratum R through Stratum Z (from bottom to top) from three sedimentary cycles.Tab.1: Korrelation der Horizonte der einzelnen Grabungsquadrate in die stratigraphischen Einheiten Schicht R bis Schicht Z (vom Liegenden zum Hangenden) aus drei Sedimentationszyklen. C L hibernate.Th e Yarimburgaz Cave is presumably a year-round dwelling concerning bats.

Fig. 6 :
Fig. 6: Lower toothrow of Rhinolophus euryale and Rhinolophus blasii.In R. euryale the p1 takes less than 50% of the plane of p4, in R. blasii the proportion of the plane is greater than 50% (after S  G 1998).
Middle Pleistocene bats from the Yarimburgaz Cave in Turkish Th race(Turkey) Pleistocene bats from the Yarimburgaz Cave in Turkish Th race (Turkey) Fig. 22: Frequency distribution of the Chiroptera within the sedimentary cycle I and III based on mandibles (exception: Rhinolophus euryale).Th e MNI based on mandibles is shown in the middle of the diagram.Abb.22: Häufi gkeitsverteilung der Chiroptera in Sedimentaionszyklus I und III auf Basis der Mandibeln (Ausnahme: Rhinolophus euryale).Die MIZ, basierend auf der Zahl der Mandibeln, ist jeweils in der Mitte des Diagramms angegeben.Since all of the documented taxa are present in the extant fauna of Turkish Th race no diff erence from the present-day climate is indicated by the bat fauna.However, the rodent and lagomorph fauna indicates a dryer, more open landscape and a cooler climate than today U because this square was very shallow and soon reached bedrock so it could not be separated in stratigraphic units.It is unfortunate that most of the bat fauna was recovered from this trench, since its precise stratigraphic level is uncertain.Middle Rhinolophus ferrumequinum, R. euryale, R  mehelyi, Myotis myotis, M. blythii, M. bechsteini,  M. nattereri, M. emarginatus, group M. mystacinus-brandti-daubentoni, M. capaccinii, Plecotus  auritus, P. austriacus, and Miniopterus schreibersi.In the quantitative distribution, R. mehelyi (56%), M. blythii (25%), and M. bechsteini (9%) are predominant(fi gs.21, 22).Th e chiropterans from Yarimburgaz Cave are signifi cant as it is the richest Middle Pleistocene bat fauna of the Eastern Mediterranean found to date.Nine to eleven additional chiropteran species are recorded for that short time of this area, fi ve had been mentioned previously (S  K ).Th ey document the composition of the modern faunal assemblage.Although the bats seem to be of little stratigraphical importance they provide new ecological information, since they may indicate a much milder climate than the rodents and lagomorphs from the same layer.Th e seasonal migration common for various bats may have allowed some to populate Turkish Th race only during summer.Th us species indicating a somewhat warmer climate were preserved together with species which characterize a distinctly cooler steppe environment.Because bats are known to migrate it is postulated that some species like Rhinolophus mehelyi and Miniopterus schreibersi were only occasional residents during that period.