MYOTIS DASYCNEME IN A LATEST PLEISTOCENE BAT

Geologica Romana 41 (2008), 117-123
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, [email protected]
**Dipartimento di Scienze Geologiche, Università di Roma Tre and Centro di Ecologia Evolutiva, L.go S. Leonardo Murialdo 1,
00146 Roma, [email protected]
***S.G.A., Via F. Rismondo 19, 05100 Terni, [email protected]
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 fredda 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 distribuzione più a sud.
PAROLE CHIAVE: Chiroptera, tardo Pleistocene, Italia centrale.
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 fractured 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 characterized 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).
118
Geologica Romana 41 (2008), 117-123
ARGENTI et al.
ing waters and the deposition of the sediments into a
small basin. The remains were founded without concretions 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,
Myotis myotis, M. bechsteini and M. dasycneme. The
only non-chiropteran remains (very fragmentary) belong
to a small mustelid of the size of a stoat (Mustela
erminea).
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 temperate Eurasia from Britain to Japan, south to north-western 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
Rhinolophus
hipposideros
L tot
L palate
W of the rostrum between C
W of the rostrum between M2
W intraorbital constriction
L P4 right
W P4 right
L M1 right
W M1 right
L M2 right
W M2 right
L P4 left
W P4 left
L M1 left
W M1 left
L M2 left
W M2 left
L M3 left
W M3 left
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 reported 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;
Dimitrijević, 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 winter (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.
1024/a 1024/b 1024/c 1024/d
13.70
3.38
3.40
4.78
2.31
0.81
1.01
1.23
1.09
1.21
1.19
0.88
0.90
1.22
1.01
1.20
1.19
13.40
3.70
3.97
4.92
1.95
1.40
0.60
0.95
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).
MYOTIS DASYCNEME IN A LATEST PLEISTOCENE BAT...
The morphology and dimensions (Tabs. 1 and 3) of the
specimens are identical with those of recent Rhinolophus
hipposideros. The small horseshoe bat is the most abundant 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 typical 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
Geologica Romana 41 (2008), 117-123
119
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 localities (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
1026
L symphysis-condyle
L symphysis-angular process
L C-M3
L M1-M3
L M1
W1 M1
W2 M1
L M2
W1 M2
W2 M2
L M3
W1 M3
W2 M3
17.20
16.90
9.59
5.39
2.08
1.22
1.46
2.10
1.35
1.45
1.87
1.30
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 = larghezza; W1 = larghezza del trigonide; W2 = larghezza del talonide;
le misure sono state effettuate secondo Storch (1974).
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).
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 (larghezza = 3, 15 mm); b) Myotis dasycneme (Boie) - Epifisi distale di
un omero sinistro (larghezza = 2,95 mm).
120
Geologica Romana 41 (2008), 117-123
Species
Rhinolophus
ferrumequinum
R. hipposideros
ARGENTI et al.
Sample
L
W caput
1023
1025/a
1025/b
1025/c
1025/d
1025/e
1025/f
1025/g
1025/h
1025/i
1025/j
1025/k
1025/l
-21.80
23.30
24.50
23.70
22.70
23.80
24.50
21.80
23.50
23.40
21.50
23.50
5.28
3.30
3.36
3.30
3.36
3.34
3.40
3.37
3.28
3.25
3.31
3.34
3.29
Myotis myotis
1027/a
1027/b
33.20
33.20
4.10
4.20
Myotis bechsteini
Myotis dasycneme
1028
1029
-23.70
3.15
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.
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 subspecies, Myotis bechsteini intermedius Rybar, was erected 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 signaled 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 (Horàček, 1976; Topál,
1981; Wołoszyn, 1987, 1989 with bibliography; Horàček
& Hanák, 1989; Roer, 2001; Ochman & Wołoszyn,
2003). Cittareale cave is the southernmost site in the past
(and present - see Tvrtkovič 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 improbable 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 individual 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
Geologica Romana 41 (2008), 117-123
MYOTIS DASYCNEME IN A LATEST PLEISTOCENE BAT...
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.
121
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 systematics. We thank also the reviewers of the manuscript Proff. M.R.
Palombo and C. Petronio.
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Accettato per la stampa: Ottobre 2008