Myotis Dasycneme in a latest Pleistocene bat assemblage of Cittareale cave (Rieti, Latium, Central Italy)

MYOTIS DASYCNEME IN A LATEST PLEISTOCENE BAT ASSEMBLAGE OF CITTAREALE CAVE (RIETI, LATIUM, CENTRAL ITALY)

Patrizia Argenti*, Tassos Kotsakis ** & Federico Sabatini***

*Dipartimento di Scienze della Terra, Università di Perugia, P.zza Università, 06100 Perugia, biogeo@unipg.it

**Dipartimento di Scienze Geologiche, Università di Roma Tre and Centro di Ecologia Evolutiva, L.go S. Leonardo Murialdo 1, 00146 Roma, kotsakis@uniroma3.it

***S.G.A., Via F. Rismondo 19, 05100 Terni, geoass@tin.it

ABSTRACT - Five species of fossil bats have been discovered in Cittareale cave (Rieti, Latium, Central Italy):

Rhinolophus ferrumequinum, R. hipposideros, Myotis (Myotis) myotis, M. (M.) bechsteini and M. (Leuconoë) dasycneme. The presence of a northern “cold” species like M. dasycneme indicates a latest Pleistocene cold phase, probably the Younger Dryas. Cittareale cave is the southernmost site of the past distribution of M. (L.) dasycneme in Europe, which at the present is found as south as northern Croatia.

KEY WORDS: Chiroptera, latest Pleistocene, Central Italy.

RIASSUNTO - Cinque specie di pipistrelli fossili sono state scoperte nella Grotta di Cittàreale (Rieti, Lazio, Italia centrale):Rhinolophus ferrumequinum, R. hipposideros, Myotis (Myotis) myotis, M. (M.) bechsteini e M.

(Leuconoë) dasycneme. La presenza di M. dasycneme, specie settentrionale “fredda”, indicherebbe una fase fred- da del Pleistocene terminale, probabilmente il Dryas Recente. La Grotta di Cittareale è la località più meridionale della distribuzione passata di tale specie, che attualmente ha nella Croazia settentrionale il suo punto di dis- tribuzione più a sud.

PAROLE CHIAVE: Chiroptera, tardo Pleistocene, Italia centrale.

Geologica Romana41 (2008), 117-123

INTRODUCTION - GEOLOGICAL SETTING

Cittàreale cave (RI) (Fig.1), open in San Rufo Valley, in the west side of Monte Pozzoni (or Monte Pizzuto), at about 1420 m a.s.l., is known since the ’sixties, but in the

’eighties explorations of GGP (Gruppo Grotte Pipistrelli) CAI Terni allowed the knowledge of the karst system (Gatti & Uffreduzzi, 1989). Along the slope, there are hypogean karst marks such as cylindric wells with a diameter of some metres, deep about ten metres and other small caves; in some of these it is possible to penetrate for a few distance (Sabatini & Gatti, 1989). The cave is set up in the marly limestone of the Scaglia Rossa, very frac- tured and faulted, due to the nearness of the thrust faults of the M. Sibillini Unit on the Gran Sasso - Cittareale Unit, which has various thrusts (thrust of Monte Prato and of Monte Pizzuto’ one) (Calamita et al., 1995). The karst cave, about 3 Km long and 450 m deep, is charac- terized by a thick interlacing of underground conduits and tunnels, situated on various levels linked by circular wells (deep up to 160 m). Particularly the main branch, which brought to the present end of the cave, is set up on a fracture system proceeding N 70° W and deeping to NE. The secondary branch is set up on a fracture system striking NNE-SSW (Preziosi & Scipioni, 1993).

The palaeontological remains was discovered in 1988 by one of us (F. S.), during the exploration which brought to the end of the cave. The fossils was found at a deep of about 100 m, about 400 m from the entrance, after many underground conduits and some wells. The fossiliferous site is in the final portion of a subhorizontal conduit transversal to a well, about 10 m long. In this

portion the underground conduit is about 1 m high and the filling is closed by a white concretional deposit. The fossil remains were found on the floor of a level, few centimetres thick, of greysh unconsolidated fine grained sediments, sands and silts, overlaining the concretional fillings. This deposit testifies an active phase of the underground passage after the filling itself, with circulat-

Fig. 1 - Location of the Cittareale cave (Rieti, Central Italy).

- Posizione geografica della Grotta di Cittareale (Rieti, Italia centrale).

ing waters and the deposition of the sediments into a small basin. The remains were founded without concre- tions on the surface of the deposits.

PALAEONTOLOGY

The fossil remains of Grotta di Cittareale belong, with few exceptions, to the order Chiroptera. Five species are present: Rhinolophus ferrumequinum, R. hipposideros,

only non-chiropteran remains (very fragmentary) belong to a small mustelid of the size of a stoat (Mustela

Systematics

Fam. Rhinolophidae Gray, 1825 Gen. Rhinolophus Lacépède, 1799

Rhinolophus ferrumequinum (Schreber, 1774)

Material: one distal epiphysis of a left humerus (n° 1023, Palaeontological Collection of Dipartimento di Scienze Geologiche, Università Roma Tre).

Locality: Grotta di Cittareale (Rieti, Latium, Central Italy).

Age: (?) latest Pleistocene.

Both morphology and dimensions (Tab. 3) of the distal epiphysis of this humerus agree very well with those of living R. ferrumequinum (see Felten et al., 1973).

The big horseshoe bat is widely distributed across tem- perate Eurasia from Britain to Japan, south to north-west- ern Africa, Palestine and northern India (Csorba et al., 2003).

During Late Pleistocene and Holocene this species was very common in many Italian sites both in mainland

and on the islands (De Gregorio, 1894; Del Campana, 1914; De Stefani, 1916; Pasa, 1953; Leonardi et al., 1962; Bartolomei, 1966; Tozzi in Cigna, 1967; Tozzi, 1970; Ponticelli & Simonetta, 1979; Sondaar et al., 1984; Capasso Barbato & Kotsakis, 1986; Bressan, 1987; Kotsakis, 1987, 1991; Esu et al., 1990; Bon &

Boscato, 1995; Alhaique et al., 1996; Taliana et al., 1996; Tata & Kotsakis, 2005; Salari, in press; Salari &

Di Canzio, in press; Tang & Kotsakis, 2008). Remains of this bat collected in sites assigned to the Middle Pleistocene are also known in Italy (Ambrosetti et al., 1979; Kotsakis & Petronio, 1980). The species is report- ed from Pirro Nord (Apulia, Italy), a fossiliferous site of Early Pleistocene age (Tata & Kotsakis, in press; Salari et al., in press). Fossils of R. ferrumequinum are signaled in many European sites and also from some localities of Asia (see Capasso Barbato & Kotsakis, 1986; Sevilla García, 1988; and references in those papers;

Dimitrijevi ć, 1997; Montoya et al., 1999, 2001; Jin et al., 2000; Popov, 2000; Alcover, 2003; Rossina et al., 2006;

Tong et al., 2008; Tata & Kotsakis, in press).

R. ferrumequinum in Europe favours woodlands and

wooded country and is common in caves during the win- ter (Csorba et al., 2003).

Rhinolophus hipposideros (Bechstein, 1800)

Material: two skulls (one without teeth) and a fragment of a third skull; twelve humeri (four right and eight left), and a distal epiphysis of a left humerus (n° 1024 (skulls) and n° 1025 (humeri), Palaeontological Collection of Dipartimento di Scienze Geologiche, Università Roma Tre).

Locality: Grotta di Cittareale (Rieti, Latium, Central Italy).

Age: (?) latest Pleistocene.

Rhinolophus

hipposideros

L tot 13.70 13.40

L palate 3.38 3.70

W of the rostrum between C 3.40 3.97 W of the rostrum between M2 4.78 4.92 W intraorbital constriction 2.31 1.95

L P4 right 0.81 1.40

W P4 right 1.01 0.60

L M1 right 1.23

W M1 right 1.09

L M2 right 1.21

W M2 right 1.19

L P4 left 0.88

W P4 left 0.90

L M1 left 1.22

W M1 left 1.01

L M2 left 1.20

W M2 left 1.19

L M3 left 0.95

W M3 left 1.00

Tab. 1 - Cranial and dental measures (in mm) of Rhinolophus hipposideros. L = length; W = width; the measures are taken following Storch (1974).

- Misure craniche e dentarie (in mm) di Rhinolophus hipposideros. L = lunghezza; W = larghezza; le misure sono state effettuate secondo Storch (1974).

Geologica Romana41 (2008), 117-123

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MYOTIS DASYCNEMEIN A LATEST PLEISTOCENE BAT...

The morphology and dimensions (Tabs. 1 and 3) of the specimens are identical with those of recent Rhinolophus

dant species in Cittareale cave.

R. hipposideros is widely distributed in the western

Palearctic from Ireland to southern Kazakhstan and Kashmir, south to north-western Africa and through western Anatolia to eastern Sudan and Ethiopia (Csorba et al., 2003). In Italian sites the small horseshoe bat is less common as fossil than the big horseshoe bat. It is reported from some Middle and Late Pleistocene sites of the mainland (Pasa, 1953; Bartolomei, 1962; Tozzi, 1969; Sala, 1973) and also from the islands (Malta, Sicily, Sardinia) (Storch, 1970, 1974; Kotsakis &

Petronio, 1980; Kotsakis, 1987). R. hipposideros is reported from a few Early Pleistocene European sites and is more common in the Middle to Late Pleistocene and Holocene deposits (see Sevilla García, 1988 with bibliography, 1990; Toskan, 2002; Alcover, 2003;

Ochman, 2003).

Like R. ferrumequinum the small horseshoe bat is common in wooded areas and during the winter is found in caves (Csorba et al., 2003).

Fam. Vespertilionidae Miller, 1897 Gen. Myotis Kaup 1829

Subgen. Myotis Kaup 1829

Myotis (Myotis) myotis (Borkhausen, 1797)

Material: a right mandibular ramus with M1-M3 and two humeri (n° 1026 (mandible) and n.° 1027 (humeri), Palaeontological Collection of Dipartimento di Scienze Geologiche, Università Roma Tre).

Locality: Grotta di Cittareale (Rieti, Latium, Central Italy).

Age: (?) latest Pleistocene.

The morphology and dimensions of hemimandible, teeth and distal epiphysis of humeri (Tabs. 2 and 3) make possible to ascribe the remains to the group of “big”

Myotis, Myotis myotis (Borkhausen) and Myotis blythii

(Tomes). After Topál & Tusnádi (1963) and Sevilla García & López Martínez (1986) the relative length of talonid and trigonid of M3 is the only one character that permits to separate the two species. In our specimen the trigonid is longer than the talonid. This character is typ- ical of M. myotis. For this reason, beside the fact that the dimensions of the teeth are rather small we identify the fossils as M. myotis.

The species is widespread in Europe ranging from England and Portugal to Ukraina and Israel. Is is also recorded from many Mediterranean islands and from the Azores. The species is common in the Late Pleistocene and Holocene deposits of Italian peninsula (Simonelli, 1917; Fabiani, 1919; Pasa, 1953; Radmilli, 1955;

Capasso Barbato & Kotsakis, 1986; Sala, 1990;

Kotsakis, 1991; Bon & Boscato, 1995; Tata & Kotsakis, 2005) and is also present in the Late Pleistocene of

Sardinia (Kotsakis, 1987). Very likely, during the cold phases of Middle and Late Pleistocene, Italy was a refuge area for this species (Ruedi et al., 2008). In Europe it is known from many Late Pleistocene sites and it is also reported from a few Middle Pleistocene locali- ties (Capasso Barbato & Kotsakis, 1986; Sevilla García, 1988 and references in those papers; Alcover, 2003).

The mouse-eared bat is common in open and slightly wooded areas and is found in caves during the whole year (Güttinger et al., 2001).

Myotis (Myotis) bechsteini (Kuhl, 1818)

Material: a distal epiphysis of a right humerus (n° 1028, Palaeontological Collection of Dipartimento di Scienze Geologiche, Università Roma Tre) (Fig. 2a).

Myotis myotis

L symphysis-condyle 17.20 L symphysis-angular process 16.90

L C-M3 9.59

L M1-M3 5.39

L M1 2.08

W1 M1 1.22

W2 M1 1.46

L M2 2.10

W1 M2 1.35

W2 M2 1.45

L M3 1.87

W1 M3 1.30

W2 M3 0.95

Tab. 2 - Dental measures (in mm) of Myotis myotis. L = length; W = width; W1 = width of the trigonid; W2 = width of the talonid; the measures are taken following Storch (1974).

- Misure dentarie (in mm) di Myotis myotis. L = lunghezza; W = lar- ghezza; W1 = larghezza del trigonide; W2 = larghezza del talonide;

le misure sono state effettuate secondo Storch (1974).

Fig. 2 - a) Myotis bechsteini (Kuhl) - Distal epiphysis of right humerus (width = 3.15 mm); b) Myotis dasycneme (Boie) - Distal epiphysis of left humerus (width = 2.95 mm).

- a) Myotis bechsteini (Kuhl) - Epifisi distale di un omero destro (lar- ghezza = 3, 15 mm); b)Myotis dasycneme (Boie) - Epifisi distale di un omero sinistro (larghezza = 2,95 mm).

Locality: Grotta di Cittareale (Rieti, Latium, Central Italy).

Age: (?) latest Pleistocene.

The distal epiphysis of a single humerus correspond both morphologically and dimensionally (Tab. 3) to the same bone of the living Myotis bechsteini (see Felten et al., 1973).

M. bechsteini

is widespread through western Eurasia from Sweden, England and Portugal to Iran (Koopman, 1994). In Italy this species is not very common and is very rare in the southern half of the Peninsula and in Sicily (Vergari et al., 1998; Agnelli et al., 2004).

As fossil M. bechsteini is not very common in Italian sites. It is known in Holocene deposits of Grotta del Lago (Triponzo, Umbria) (Taliana et al., 1996), in Late Pleistocene deposits of Buco dell’Orso cave (Como, Lombardy) (Santi, 2000), of Broion cave (Veneto) (Pasa, 1953), of M. Cucco cave (Umbria) (Capasso Barbato &

Kotsakis, 1986), of the fissure “XI-dic. 2001” of Monte Tuttavista (Sardinia) (Abbazzi et al., 2004), and at Spinagallo (Syracuse, Sicily), in a deposit of Middle Pleistocene age (Kotsakis & Petronio, 1980).

Storch (1974), assigned some remains from Ghar Dalam cave (Malta) of Early- Middle Pleistocene age to an extinct subspecies, Myotis bechsteini robustus Topal, errected on material of Early Pleistocene age from Hungary (Topál, 1963), whilst another extinct sub- species, Myotis bechsteini intermedius Rybar, was erect- ed for some Holocene remains from former Czechoslovakia (Rybar, 1976). Bechstein’s bat is rather common in European sites of Late Pleistocene (and Holocene) age and is known also from some localities of

Middle and Early Pleistocene age. It’s appearance go back to the Pliocene (Capasso Barbato & Kotsakis, 1986 with bibliography; Wołoszyn, 1987, 1989; Sevilla García, 1988 with bibliography, 1990; Aguilar, 1998;

Baagøe, 2001; Toskan, 2002; Ochman, 2003; Ochman &

Wołoszyn, 2003; Blant et al., 2007). After Topál (1985) the Pliocene species Myotis gundersheimensis Heller, shows close affinities with M. bechsteini.

As the two horseshoe bats M. bechsteini is common in wooded areas and during the winter is found in caves (Baagøe, 2001).

Subgen. Leuconoë Boie, 1830

Myotis (Leuconoë) dasycneme (Boie, 1825)

Material: a left humerus (n° 1029, n° Palaeontological Collection of Dipartimento di Scienze Geologiche, Università Roma Tre) (Fig. 3).

Locality: Cittareale cave (Rieti, Latium, Central Italy).

Age: (?) latest Pleistocene.

This single left humerus is identical morphologically to the correspondent bone of living Myotis dasycneme (see Felten et al., 1973).

The pond bat ranges from northwestern Europe east to central Siberia and northeastern China (Koopman, 1994). In Italy only one erratic individual has been sig- naled in 1881 in Veneto (Lanza & Agnelli, 1999).

M. dasycneme

is present in a few Late Pleistocene Italian sites: Broion cave (Pasa, 1953), Mezzena shelter (Veneto) (Sala, 1990), Grotta del Vento (Iesi, Marche) (Esu et al., 1990). It is signaled in some Pleistocene and Holocene sites of central Europe (Horà ček, 1976; Topál, 1981; Wołoszyn, 1987, 1989 with bibliography; Horà ček

& Hanák, 1989; Roer, 2001; Ochman & Wołoszyn, 2003). Cittareale cave is the southernmost site in the past (and present - see Tvrtkovi č et al., 2001) distribution of the species in Europe.

Rather rare in the living faunas, it dwells in wooded areas near ponds. During the winter it is found in caves (Roer, 2001).

CONCLUSIONS

The remains of the assemblage are collected in surface but the bones are fossilized and clearly differ from the remains of recent bats. An attribution to a precise time interval is impossible; nevertheless it is highly improba- ble the conservation of such delicate bones like those of the bats for long periods exposed on the surface, even in protected areas like the interior part of the caves. The presence of a northern species, M. dasycneme, known in Italy only in cold fossil assemblages and absent today from the mammalian fauna of the region (only one indi- vidual has been signaled in the north-eastern Italy during the XIX century), suggests to attribute the assemblage of Cittareale cave to the last cold interval. The Younger

Species Sample L W caput

Rhinolophus

ferrumequinum 1023 -- 5.28

R. hipposideros 1025/a 21.80 3.30

1025/b 23.30 3.36

1025/c 24.50 3.30

1025/d 23.70 3.36

1025/e 22.70 3.34

1025/f 23.80 3.40

1025/g 24.50 3.37

1025/h 21.80 3.28

1025/i 23.50 3.25

1025/j 23.40 3.31

1025/k 21.50 3.34

1025/l 23.50 3.29

Myotis myotis 1027/a 33.20 4.10

1027/b 33.20 4.20

Myotis bechsteini 1028 -- 3.15

Myotis dasycneme 1029 23.70 2.90

Tab. 3 - Measures of the humeri (in mm). L = total length of humerus;

W = width of distal epiphysis.

- Misure degli omeri (in mm). L = lunghezza totale dell’omero; W = larghezza dell’epifisi distale.

MYOTIS DASYCNEMEIN A LATEST PLEISTOCENE BAT... Geologica Romana41 (2008), 117-123

121 Dryas period (11.500 - 12.650 BP - Ravazzi et al., 2007)

is the most probable one. We suggest this period because is the last temporal interval with low temperatures in the latitude of the cave. Obviously the age of the association is inferred and a younger (for example cold oscillations strongly affecting the distribution of many species are known also in very recent Holocene times - Esu &

Girotti, 1991 for non-marine molluscs in central Italy) or perhaps older age is possible even rather improbable.

The five species collected in Cittareale cave indicate a wooded environment with open space and ponds.

AKNOWLEDGEMENTS - The Authors thank Gruppo Grotte Pipistrelli CAI Terni for the collection of the material and Dr. L. Salari for the usufull discussions on bats systemat- ics. We thank also the reviewers of the manuscript Proff. M.R.

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Accettato per la stampa: Ottobre 2008