SIPHONAPTERA OF SQUIRRELS AND DORMICE (RODENTIA: SCIURI-DAE, GLIRIDAE) FROM THE WESTERN AND CENTRAL BALKANS
Savo BRELIH and Tomi TRILAR
Slovenian Museum of Natural History, Prešernova 20, P.O.Box 290, SI-1001 Ljubljana, Slovenia, e-mail: ttrilar@pms-lj.si
Abstract - We present data on 22 species and 6 subspecies of Siphonaptera of Sciurus vulgaris, Spermophilus citellus, Glis glis, Diyomys nileclula, Eli o my s quercinus, and Muscardinus avellanarius from the Western and Central Balkans. A new subspecies is described: Ctenophthalmus orientalis jakupicae ssp. n. The distribution of 1 species and 8 subspecies of fleas typical for the mentioned hosts is shown on 6 maps. A survey of hosts is presented as well as the Siphonaptera on them. Postglacial migrations of S. citellus are suggested on the basis of their variability as well as the distribution of the Siphonaptera.
Key words: Siphonaptera, Sciurus vulgaris, Spermophilus citellus, Glis glis, Diyomys nit-edula, Eliomys quercinus, Muscardinus avellanarius, Balkans
Izvleček - BOLHE VEVERIC IN POLHOV (RODENTIA: SCIURIDAE, GLIRIDAE) ZAHODNEGA IN OSREDNJEGA BALKANSKEGA POLOTOKA
Navedenih je 22 vrst in 6 podvrst bolh z veverice (Sciurus vulgaris), tekunice (Spermophilus citellus), navadnega polha (Glis glis), drevesnega polha (Diyomys nitedu-la), vrtnega polha (Eliomys quercinus) in podleska (Muscardinus avellanarius) iz zahodnega in osrednjega Balkanskega polotoka. Opisana je nova podvrsta, Ctenophthalmus orientalis jakupicae ssp. n. Razširjenost ene vrste in 8 podvrst bolh značilnih za navedene gostitelje je prikazana na 6 geografskih kartah. Podan je tudi pregled gostiteljev in na njih ugotovljenih bolh. Na osnovi razširjenosti bolh in variabilnosti tekunic je predlagana shema postglacialnih selitev tekunice.
Ključne besede: bolhe, veverica, tekunica, navadni polh, drevesni polh, vrtni polh, podlesek, Balkanski polotok
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Introduction
This contribution continues the series »Ectoparasitical entomofauna of Yugoslav mammals« (Brelih & Petrov, 1987; Brelih, 1986; Brelih & Trilar, 2000). The changes in former Yugoslavia in the last few years resulted in the establishment of five new countries and thus the title of the series is now inappropriate (Fig. 1). We will call this area as the Western and Central Balkans.
The present paper provides the data on siphonapterofauna of squirrels (Sciuridae) and dormice (Gliridae) from this territory.
Wagner (1928-1929, 1934, 1936, 1939), Rosicky & Carnelutti (1959), Rosicky & Todorovič (1964), Hopkins & Rothschild (1962, 1966, 1971), Brelih (1986) and Trilar (1995, 1997) published data on fleas of squirrels (Sciuridae) and dormice (Gliridae) from the territory of the Western and Central Balkans.
Data concerning hosts are summarized from Dulič & Mirič (1967), Ružič (1978), Petrov (1992), Krystufek (1991, 1993, 1996), Krystufek & Hrabe (1996) and Tvrtkovič et al. (1995). We used the mammalian nomenclature according to Mitchell-Jones et al. (1999).
Material and methods
Most of the material was collected in the last 30 years along with the mammalian field collecting. There were no field trips purely for ectoparasite collection. The only exception was the systematic study of Glis glis nests in nestboxes at the Mt. Snežnik and Mt. Goteniška Gora (Trilar, 1995, 1997). Dormice were mainly collected in micro-mammalian faunistic studies, where between 100 and 200 snap-traps were placed per night. Dead animals were removed from the traps early in the morning as numerous ectoparasites later leave them. Each single species of host was stored into a separate linen or plastic bag until the examination attended to immediately after collection. Ectoparasites were stored in 70% alcohol, separately with respect to host, survey site, altitude, and date. In most cases ecological examinations were not performed. Unfortunately, we very rarely possessed information on the number of host specimens examined. European sousliks (Spermophilus citellus) were collected in all distribution areas, but in limited numbers. Red squirrels (Sciurus vulgaris) were mainly collected by shooting and also recovered as traffic casualties. We examined material provided by hunters, where animals had been dead for a longer time and some fleas had already left them. A small number of S. vulgaris were inspected from Slovenia, but only single specimens from all other areas. Until today we have not examined a single specimen of the Alpine marmot (Marmota marmota). Collected flea specimens were used to prepare microscopic slides in Canada balsam, with the exception of some large samples of Ceratophyllus sciurorum sciurorum from nests of Glis glis, which were preserved in 70% alcohol. All the material without a literature citation is kept in the Slovenian Museum of Natural History (coll. S. Brelih, coll. T. Trilar). Species identification was according to Hopkins & Rothschild (1953, 1956, 1962, 1966, 1971) and Rosicky (1957). Some of the more recent contributions of authors (Cyprich, 1989) were also utilized.
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List of the hosts: Parus major Linnaeus, 1758; Talpa europaea Linnaeus, 1758; Homo sapiens Linnaeus, 1758; Sciurus vulgaris Linnaeus, 1758; Marmota marmota (Linnaeus, 1758); Spermophilus citellus (Linnaeus, 1766); Spermophilus suslicus (Giildenstaedt, 1770); Spermophilus pygmaeus (Pallas, 1779); Cricetus cricetus (Linnaeus, 1758); Clethrionomys glareolus (Schreber, 1780); Dinaromys bogdanovi (Martino, 1922); Microtus aivalis (Pallas, 1778); Nannospalax leucodon (Nordmann, 1840); Micromys minutus (Pallas, 1771); Apodemus flavicollis (Melchior, 1834); Apodemus mystacinus epimelas (Nehring, 1902); Apodemus sylvaticus (Linnaeus, 1758); Rattus rattus (Linnaeus, 1758); Mus musculus s. lat. Linnaeus, 1758; Glis glis (Linnaeus, 1766); Muscardinus avellanarius (Linnaeus, 1758); Eliomys quercinus (Linnaeus, 1766); Diyomys nitedula (Pallas, 1778); Vulpes vulpes (Linnaeus, 1758); Martes foina (Erxleben, 1777); Martes martes (Linnaeus, 1758); Meles meles (Linnaeus, 1758); Felis silvestris Schreber, 1775.
Fig. 1: Western and Central Balkans with indicated states, borders, and UTM (100 x 100 km grid).
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Fig. 2: Map of the Western and Central Balkans. Numbers indicate survey sites (localities).
SLOVENIJA (SLOVENIA):
1.	VL07 Komen, Kobjeglava
2.	VL08 Trnovski gozd: Caven
3.	VL38 Hotedršica, Novi Svet; Hotedršica, Ravnik
4.	VL54 Snežnik; Snežnik: Sviščaki
5.	VL58 Borovnica, Pako
6.	VL68 Turjak
7.	VL75 Goteniška gora: Draga
8.	VL85 Kočevje, environs
9.	VM02 Bohinj: Ukane
10.	VM04 Trenta: Julijami
11.	VM05 Rateče, Zelenci
12.	VM21 Cerkno
13.	VM30 Poljanska dolina
14.	VM32 Jelovica: Goška ravan; Rudno; Rudno, Hujska
15.	VM42 Kranj, Sveti Jošt
16.	VM43 Tržič
17.	VM44 Podljubelj, Čižovnik
18.	VM50 Ljubljana: Vrhovci
19.	VM51 Medvode; Sorsko polje: Žabnica
20.	VM52 Cerklje, Štefanja gora; Kranj; Kranj, Milje; Šenčur, Voglje
21.	VM53 Lom, Grahovše, Gaberčev rovt
22.	VM60 Črnuče; Ljubljana and environs; Gameljne
23.	VM61 Kamnik, environs
24.	VM63 Kamniška Bistrica
25.	VM64 Zgornje Jezersko
26.	VM80 Litija
27.	WL08 Mirna na Dolenjskem
28.	WL27 Kostanjevica, Šentjernej
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29.	WM05 Slovenj Gradec
30.	WM34 Pohorje: Lobniai
31.	WM40 Kozje, Lesično, Gruska jama
32.	WM43 Poljčane, Studenice, Jama v kamnolomu
33.	WM65 Dvorjane, Vurberg, Grmada
34.	WM95 Srednja Bistrica HRVATSKA (CROATIA):
35.	VK78 Krk Island: Veli vrh
36.	VK95 Velebit: Mirevo
37.	VK96 Velebit: Zavižan; Zavižan, Modrica dolac (Botanički vrt); Vučjak
38.	VL73 Gorski Kotar: Risnjak
39.	WK01 Pag Island: Pag, environs
40.	WK02 Pag Island: Metajna
41.	WK13 Velebit: Baske Oštarije; Papratnjak
42.	WK20 Ražanac, Vrankoviči
43.	WK41 Velebit: Bunjevac
44.	WK50 Velebit: Tulove grede
45.	WK60 Velebit: Predzid
46.	WL84 Vukomeričke gorice: Prkovec
47.	WL95 Turopolje: Peščenica, near the river Odra
48.	WL97 Zagreb, Dugo Selo
49.	XH28 Hvar Island: Brusje 50 XH38 Brač Island: Dol
51.	XJ11 Split
52.	XJ60 Biokovo: Sveti Jura
53.	BN62 Dubrovnik
BOSNA I HERCEGOVINA (BOSNIA AND HERZEGOVINA):
54.	XJ29 Šator planina: Salonsko jezero; Babina greda
55.	XJ93 Svinjača, Blidinje jezero
56.	YJ31 Velež, Rujište
57.	BN88 Bjelašnica (Gacko)
58.	BP84 Igman
59.	CN19 Maglič
60.	CP00 Zelengora: Orlovačko jezero
61.	CP26 Gazivoda
JUGOSLAVIJA: ČRNA GORA (YUGOSLAVIA: MONTENEGRO):
62.	BN90 Hercegnovi
63.	CM29 Cetinje. environs
64.	CN02 Orjen
65.	CN94 Bjelasica: planina Jelovica; Zekova Glava JUGOSLAVIJA: SRBIJA: VOJVODINA (YUGOSLAVIA: SERBIA: VOJVODINA):
66.	CQ79 Srem: Erdevik
67.	CR84 Bačka: Savino Selo
68.	DQ09 Srem: Fruška Gora: Irig; Jazak
69.	DQ78 Pančevo, Crepaja
70.	DR00 Srem: Fruška Gora: Zmajevac
71.	DR20 Srem: Fruška Gora: Čortanovci
72.	DR30 Bačka: Titel, Lok
73.	DR44 Banat: Melenci
74.	DR61 Banat: Zrenjanin, Orlovat, reka Tamiš
75.	DR80 Deliblatska Peščara: Samoš
76.	EQ06 Deliblatska Peščara: Deliblato; Dolina
77.	EQ08 Deliblatska Peščara: Čoka
78.	EQ16 Deliblatska Peščara: Šumarak
79.	EQ17 Deliblatska Peščara: Kremenjak; Šušara
80.	EQ26 Banat: Banatska Palanka JUGOSLAVIJA: SRBIJA (YUGOSLAVIA: SERBIA):
81.	CP66 Tara Planina: Kremiči; Predov Krst
82.	DP50 Golija: Biser Voda
83.	DQ60 Arandelovac
84.	EN89 Niš: Sičevo
85.	EN98 Suva Planina
86.	EP30 Veliki Jastrebac, Ravnište
87.	EQ82 Donji Milanovac
88.	FN00 Besna Kobila
89.	FN02 Vlasinska Tresava
90.	FN03 VI asi na
91.	FN08 Suva Planina: Tri lokve
92.	FN38 Vidlič: Basara: Planinica
93.	FN48 Stara Planina: Dojkinci; Dojkinci, Bašta; Ponor
94.	FN49 Stara Planina; Stara Planina: ICopren JUGOSLAVIJA: SRBIJA: KOSOVO (YUGOSLAVIA: SERBIA: KOSOVO):
95.	DN22 Rugovo, Bjeluha; Kučište
96.	DN33 Žljeb: Kula
97.	DN53 Istok
MAKEDONIJA (MACEDONIA):
98.	DM60 Šar Planina: Lukovo Pole
99.	DM62 ICorab: Guri Velpnis
100.	DM70 Bistra
101.	DM72 Ničpurska Planina: Ničpur
102.	DM90 Kičevo, Dolenci
103.	DM95 Šar Planina: Jelak
104.	EL47 Prilep
105.	EM22 Karadžica
106.	EM3 I Jakupica: Gorno Begovo, Begovo Pole; Solunsko Pole
107.	FL06 Kožuf
108.	FL25 Gevgelija
109.	FL46 Dojl ansko jezero, Ačikot
The UTM coordinates are followed by the major localities (province, mountain, etc.), then the exact survey site. If separated by a comma, the exact locality is situated close to the major one (e.g. Kicevo, Dolenci). Tf separated by a colon, the survey site is situated at the major locality or represents its component (e.g. Sar Planina: Jelak).
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List of the species and subspecies
PULICIDAE
1.	Echidnophaga murina (Tiraboschi, 1903) Hrvatska:
29 Hvar Island: Brusje, 340 m, 4. 10. 1975, from Eliomys quercinus, leg. N. Tvrtkovič
The major host of E. murina are rats, especially R. rattus. Only rarely is found to infect other species of small ground rodents. This uncommon flea species (Hopkins & Rothschild, 1953) is distributed in the Mediterranean area (Italy, Greece, Egypt, Maroco, Turkey, Asia Minor) but was also found in Gruzia ( = Georgia) and eastern Asia (Hong Kong, Japan). The island of Hvar is the only known survey site for this area. Eliomys quercinus is the first known dormice host of this species.
2.	Pulex irritans Linnaeus, 1758 SI oven ij a:
l9 Hotedršica, Ravnik, 550 m, 9. 9. 1966, from Glis glis, leg. S. Brelih & R. Jelinčič
The passage of the human flea to G. glis is presumably a coincidental. Wagner (1939) also found P. irritans on G. glis, but the exact locality was not given.
HYSTRICHOPSYLLIDAE
3.	Hystrichopsylla (Hystrichopsylla) orientalis orientalis Smit, 1956 Bosna i Hercegovina:
l<3 Gazivoda, 26. 10. 1988, from Glis glis, leg. B. Krystufek
Hystrichopsylla o. orientalis occurs in the fur and in the nests of small terrestrial mammals (Talpidae, Soricidae, Arvicolidae and Muridae). The passage to G. glis is probably very rare.
4.	Neopsylla setosa spinea Rothschild, 1915 Jugoslavija: Srbija: Vojvodina:
2Cf 49 Srem: Fruška Gora: Jazak, 180 m, 12. 4. 1989, from Spermophilus citellus, leg. B. Krystufek & T. Trilar
Certanovci, Srem (correct Srem: Fruška Gora: Cortanovci), from Citellus citellus (=Spermophilus citellus), leg. J. Wagner (»Neopsylla setosa«) (Wagner, 1939) l9 Deliblatska Peščara: Deliblato, 100 m, 7. 5. 1947, from Spermophilus citellus, leg. A. Ružič
id 39 idem, 5. 4. 1949, leg. B. Petrov l9 Deliblatska
peščara: Čoka, 7. 5. 1947, from Citellus citellus (=Spermophilus citellus), leg. A. Ružič (»Neopsylla setosa«) (Rosicky & Todorovič, 1964) Neopsylla setosa mainly lives in the nests of Spermophilus (Hopkins & Rothschild, 1962). While we examined only the fur of collected sousliks and not the nests, the findings in the Central Balkans were quite rare. The species is probably more common than thought and probably occurs throughout the range of the host.
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Fig. 3: Distribution of Spermophilus citellus (Linnaeus) (adapted from Petrov, 1992), and survey sites of Neopsylla setosa spineci Rothschild.
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5.	Dorcitopsyllci dasycnema dasycnema (Rothschild, 1897)
»All over the territory of the Western and Central Balkans«, from Sorex and Talpa, and also from Sylvaemus flavicollis (=Apodemus flavicollis) and G lis glis (»Dorcitopsyllci dasycnemus«) (Wagner, 1939)
6.	Ctenophthalmus (Spalacoctenophthalmus) monticola (Kohaut, 1904) Jugoslavija: Srbija:
l9 Tara Planina: Kremiči, 950 m, 20. 6. 1949, from Glis glis, leg. B. Petrov 1C? Stara Planina, October 1947, from Glis glis, leg. B. Petrov & A. Ružič
The passages of fleas, typical for N. leucodon (Hopkins & Rothschild, 1962), to G. glis are presumably very rare.
7.	Ctenophthalmus (Ctenophthalmus) agyrtes wagnerianus Peus, 1950 Slovenij a:
l9 Hotedršica (=Hotedršica, Novi svet), 700 m, 26. 8. 1956, from Glis glis, leg. R.
& N. Jelinčič (»Ctenophthalmus ag)>rtes camicus«) (Rosicky & Carnelutti, 1959) l9 Hotedršica, Ravnik, 580 m, 13. 9. 1971, from Glis glis, leg. R. Jelinčič 5(3 29 Snežnik, Sviščaki, 1250 m, 4. 7. 1995, from 5 nests of Glis glis, leg. T. Trilar 1C? ibidem, 22. 7. 1997, from 1 nest of Glis glis l9 ibidem, 3. 9. 1999, from 1 nest of Glis glis
l9 Trenta: Julijana, 720 m, 13. 9. 1968, from Glis glis, leg. S. Brelih & J. Dovič
8.	Ctenophthalmus (Ctenophthalmus) agyrtes graecus Jordan, 1926 Makedonija:
l9 Prilep, 700 m, 21. 4. 1969, from Diyomys nitedula, leg. B. Petrov 1C? Jakupica: Gorno Begovo, Begovo Pole, 1950-2000 m, 11. 5. 1989, from Spennophilus citellus, leg. B. Krystufek
9.	Ctenophthalmus (Ctenophthalmus) agyrtes ohridanus Wagner, 1939 dinar us
Rostigayev, 1959 Hrvatska:
4C? 69 Brač Island: Dol, 600 m, 4. 9. 1975, from Glis glis, leg. N. Tvrtkovič
Ctenophthalmus agyrtes is one of the most common flea species on small terrestrial mammals (Talpidae, Soricidae, Arvicolinae, and Muridae). The passage to dormice is common; to sousliks is quite rare.
10.	Ctenophthalmus (Medioctenophthalmus) nifetodes brelihi Rosicky & Carnelutti, 1959 Slovenija:
3C? 39 Hotedrščica (=Hotedršica, Novi svet), 700 m (not 1264 m), 26. 8. 1956, from Glis glis, leg. R. & N. Jelinčič (Holotype and 5 paratypes) (Rosicky & Carnelutti, 1959) (included id" and l9 paratype from Rothschild collection in the British Museum (Hopkins & Rothschild, 1966)) l9 ibidem, 4. 8. 1967, from Glis glis, leg. S. Brelih & R. Jelinčič (Brelih, 1986)
The single known host of this flea subspecies is G. glis. The flea type locality is Novi
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Svet near Hotedršiea in Slovenia and, according to current knowledge, is the only known site. In Slovenia there are no known recent survey sites of D. bogdanovi, which is the major host of the remaining subspecies of Ct. nifetodes. Pleistocene fragments (D. cf. bogdanovi) are encountered in the environs of Ilirska Bistrica and the fossil Dinaromys dalmatinus has been found near Trieste in Italy (Bartolomei, 1970). D. bogdanovi is most probably extinct in Slovenia and its specific fleas presumably passed from the primary host to the new one, which is G. glis.
11.	Ctenophthalmus (Medioctenophthalmus) nifetodes tvrtkovici Brelih, 1986 Hrvatska:
l9 Velebit: Zavižan, Modrica dolac, 1460 m, 8. 9. 1979, from Glis glis, leg. N. Tvrtkovič (Brelih, 1986)
12.	Ctenophthalmus (Medioctenophthalmus) nifetodes nifetodes Wagner, 1933 Bosna i Hercegovina:
29 Bjelašnica (Gacko), 1600 m, 4. 8. 1970, from Glis glis, leg. B. Petrov (Brelih, 1986) Jugoslavija: Črna Gora:
i9 Cetinje, environs, from Glis glis, leg. V Martino (»Ctenophthalmus nivalis nifetodes« - holotype) (Wagner, 1928-29, 1933)
As many G. glis nest in rocky cracks and cavities (Polak, 1997), they come in contact with D. bogdanovi. Fleas are quite commonly pass from one host to another.
13.	Ctenophthalmus (Euctenophthalmus) congener congener Rothschild, 1907 Slovenija:
lCf Trnovski gozd: Čaven, 1240 m, 23. 9. 1971, from Glis glis, leg. S. Brelih & R. Jelinčič
14.	Ctenophthalmus (Euctenophthalmus) congener troilus Peus,1954 Makedonija:
lCf Gevgelija, 75 m, 26. 4. 1971, from Dryomys nitedula, leg. B. Petrov
Both subspecies of Ct. congener live on many species of small terrestrial mammals and periodically infect dormice.
15.	Ctenophthalmus (Euctenophthalmus) orientalis orientalis (Wagner, 1898) Jugoslavija: Srbija: Vojvodina:
ICS Torža, Bačka (correct Bačka: Savino Selo), from Crocidura sp., leg. J. Wagner
(»Ctenophthalmus orientalis«) (Wagner, 1939) Irig, Srem (Srem: Fruška Gora: Irig), from Citellus citellus (=Spermoplulus citellus),
leg. V Martino (»Ctenophthalmus orientalis«) (Wagner, 1928-29) l9 Trig, Banat (correct Srem: Fruška Gora: Irig), 27. 8. 1933, from Citellus c. citellus
(=Spermophilus citellus), leg. T. Kasapski (Hopkins & Rothschild, 1966) 3tf 39 Srem: Fruška Gora: Jazak, 180 m, 12. 4. 1989, from Spermophilus citellus, leg.
B. Kryštufek & T. Trilar lCi Crepaja, Banat (=Pančevo, Crepaja), 80 m, 17. 4. 1928, from Citellus citellus
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(=Spermophilus citellus), leg. V Martino (Hopkins & Rothschild, 1966) 3Cf 59 Banat: Banatska Palanka, 85 m, 29. 6. 1988, from Spermophilus citellus, leg. B.
Krystufek 3(3 39 idem, 30. 6. 1988
2C?Banat: Zrenjanin, Orlovat, reka Tamis, 81 m, 13. 4. 1989, from Spermophilus
citellus, leg. B. Krystufek & T. Trilar 29 Backa: Titel, Lok, 80 m, 14. 5. 1989, from Spermophilus citellus, leg. B. Krystufek Milenci, Backa (correct Banat: Melenci), from Citellus citellus (=Spermophilus citellus),
leg-V Martino (»Ctenophthalmus orientalis«) (Wagner, 1928-29) id 2V Deliblatska Pescara: Samos, 100 m, 14. 5. 1989, from Spermophilus citellus, leg. B. Krystufek
2d IV Deliblatska Pescara, 9. 12. 1981, from Talpa europciea, leg. A. Ruzic l9 Deliblatska Pescara: Deliblato, 100 m, 7. 5. 1947, from Spermophilus citellus, leg. A.
Ruzic 29 idem, 20. 4. 1949 id l9 idem, 21. 4. 1949
id Deliblatska Pescara: Dolina, 130 m, 14. 3. 1948, from Talpa europaea, leg. B. Petrov
lCf 29 Deliblatska pescara: Coka, 7. 5. 1947, from Citellus citellus (=Spermophilus
citellus), leg. A. Ruzic (»Ctenophthalmus orientalis«) (Rosicky & Todorovic, 1964) i9 Deliblatska Pescara: Sumarak, 110 m, 7. 2. 1983, from Nannospalax leucoclon, leg.
A. Ruzic 29 idem, 29. 4. 1983
l9 Deliblatska Pescara: Susara, 170 m, 7. 4. 1964, from Apodemus sylvciticus, leg. D. &
N. Heneberg Jugoslavia: S r b i j a:
id" Vlasinska Tresava, 1200 m, 18. 5. 1948, from Spermophilus citellus, leg. B. Petrov 2d 89 Vlasina, 1250 m, August 1947, from Citellus citellus (=Spermophilus citellus),
leg. A. Ruzic (»Ctenophthalmus orientalis«) (Rosicky & Todorovic, 1964) 2d 89 ibidem, 18. 8. 1947, from Talpa europaea, leg. A. Ruzic (»Ctenophthalmus orientalis«) (Rosicky & Todorovic, 1964) 60 39 ibidem, 25. 7. 1977, from Spermophilus citellus, leg. B. Petrov ici 89 Stara Planina: Ponor, 17. 6. 1947, from Citellus citellus (=Spermophilus citelus), leg. A. Ruzic (»Ctenophthalmus orientalis«) (Rosicky & Todorovic, 1964) Ctenophthalmus o. orientalis is a typical flea of souslik nests, but is frequently found also in the fur of collected animals. Usually occurs in spring and autumn, but in Yugoslavia was found also from June to August. In Yugoslavia and Macedonia it was found only within the range of its major host S. citellus. Other hosts from this area are T. europaea, Crocidura sp., N. leucodon, and A. sylvciticus.
16. Ctenophthalmus (Euctenophthalmus) orientalisjakupicae ssp. n.
(Figs. 4, 5, 7, 9) Makedonija:
some 9 Karadica (=Karadzica), from Citellus citellus (=Spermophilus citellus),
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leg. J. Wagner (»Ctenophthalmus orientalis«) (Wagner, 1939) (We did not check Wagner's material, but it probably belongs to Ct. o. jakupicae) 24Cf 299 Jakupica: Gorno Begovo, Begovo Pole, 1950-2000 m, 11. 5. 1989, from
Spermophilus citellus, leg. B. Krystufek 49 idem, 27 6. 1989
Holotype male (PMSL-IA-P-7495/1), allotype female (PMSL-IA-P-7485/2) from Begovo Pole in the Mt. Jakupica (Macedonia), 11. 5. 1989, S. c. karamani Martino & Martino, leg. B. Krystufek, paratypes of 23 males and 28 females (data as above) and 4 females from Begovo Pole in the Mt. Jakupica (Macedonia), 27. 6. 1989, from the same host, leg. B. Krystufek. The type material resides in the Slovenian Museum of Natural History in Ljubljana (PMSL - Prirodoslovni muzej Slovenije v Ljubljani, coll. S. Brelih).
Diagnosis: males of Ct. o. jakupicae differ from the nominate form in the shape of the movable process of the clasper and in the larger number of sensillae at the anterior
	No. of S3	No. of sensillae	Average
ssp. orientalis			
Vojvodina (7 populations)	15	6 - 7 (exceptionally 8)	6.87
south Serbia (Vlasina)	7	6-7	6.50
ssp. jakupicae Mt. Jakupica	24	7 - 8 (exceptionally 6)	7.42
margin of this process. Females are characterized by the apical margin of sternum VII, which forms only a very shallow sinus between the median and ventral lobes. Description: Male: For most characters Ct. o. jakupicae matches its nominate form. The most distinct character is the apical margin of the movable process of the clasper, which in the new subspecies is straight (Fig. 4, 5) while in the nominate subspecies it is concave (Fig. 6). Both sclerites which surround the apical margin in ssp. orientalis are
	No. of	No. of small bristles	No. of large bristles	Total No. of bristles
ssp. orientalis Vojvodina (7 populations) south Serbia (Vlasina)	21 2	1-8 (3.90) 6 (6.00)	5-10(5.24) 6 - 7 (6.50)	6-13(9.14) 12-13(12.50)
ssp. jakupicae Mt. Jakupica	32	3 - 9 (5.78)	5-10(6.78)	9-17(12.66)
expressed as projections which create the concavity (Fig. 6). In Ct. o. jakupicae sclerites do not go beyond the level of the anterior and apical margins (Fig. 4, 5). At the transition to posterior margin is a small projection or sclerotized tooth. The number of sensillae at the anterior margin of the movable process in the new subspecies is on average more than in the nominate form:
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Fig. 4: Clenophthalmus orientalis jakupiccie ssp. n., processes of clasper of holotype male. Arrow depicts straight apical margin.
Fig. 5: Ctenophthalmus orientalis jakupiccie ssp. n., suprafoveal portion of the movable process of four paratype males. Specimens a-d posses straight apical margin as in Fig. 4.
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S. Brel ill, T Tri lar: Siphonaptera of.squirrels and dormice
c	d
Fig. 6: Ctenophthalmus orientalis orientalis (Wagner), suprafoveal portion of the movable process of male: a. Orlovat, Banat, Vojvodina; b. Jazak, Fruška Gora, Srem, Vojvodina; c. Deliblatska Peščara, Banat, Vojvodina; d. Vlasina, southeastern Serbia. Arrow in 6 c depicts concave apical margin also seen in 6 a, b, d.
a	b
Fig. 7: Ctenophthalmus onentalis jakupicae ssp. n.: sternum VII of female: a. allotype; b. paratype. Arrow in 7 a depicts shallow sinus between the median and ventral lobes also seen in 7 b.
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Fig. 8: Ctenophlhalmus orientalis ovientalis (Wagner): sternum VII of female: a. Lok, Titel, Backa, Vojvodina; b. Jazak, Fruska Gora, Srem, Vojvodina. Arrow in 8 a depicts deeply rounded sinus between the median and ventral lobes also seen in 8 b.
One male from Vlasina (Fig. 6d) and one male from the Mt. Jakupica (Fig. 5b) are intermediate between the two subspecies in the movable process of the clasper.
Female: outline of apical margin of sternum VII forms a very shallow sinus between the median and ventral lobes (Fig. 7); in the nominate subspecies it forms a deeply rounded sinus (Fig. 8). The number of bristles on sternum VII in ssp. jcikiipiccie is on average larger than in ssp. orientalis from Vojvodina, and possibly also for the population from south Serbia (N = 2 females from Vlasina):
Bulic & Miric (1967) reported 5 subspecies of S. citellus from the Western and Central Balkans, but Krystufek (1996) questioned their validity. The subspecies S. c. karamani Martino & Martino from the Mt. Jakupica, Macedonia, is phenetically the most distinct (Krystufek, 1996). This marginal population lives in the mountains and is completely isolated. The population of Ct. orientalis collected from S. c. karamani is also very distinct from all other European populations and is described as a new subspecies.
LEPTOPSYLLIDAE
17. Leptopsylla (Leptopsylla) segnis (Schonherr, 1811) H r v a t s k a:
1CJ 29 Brae Island: Dol, 600 m, 4. 9. 1975, from Glisglis, leg. N. Tvrtkovic
Leptopsylla segnis is a cosmopolitan and sinantropic species whose major host is M. musculiis s. lat. Together with its host it frequently invades buildings inhabited by humans, especially in rural areas. But the flea can also be found far from human
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S. Brel ill, T Tri lar: Siphonaptera of.squirrels and dormice
places. From the house mouse it frequently passes to other hosts, especially insectivores and rodents, as well as to squirrels and dormice.
18.Leptopsylla	(Leptopsylla) sciurobia (Wagner, 1934) Bosna i Hercegovina:
1(3 Maglič, 1700 m, 10. 9. 1970, from Dryomys nitedula, leg. B. Petrov l9 Zelengora: Orlovačko jezero, 1500 m, 9. 9. 1984, from Dtyomys nitedula, leg. B.
Krystufek Jugoslavija: Črna Gora:
1Cs Bjelasica: planina Jelovica, 1500 m, 20. 9. 1981, from Dtyomys nitedula, leg. B.
Petrov & M. Milenkovič Jugoslavija: Srbija: Kosovo:
29 Rugovo, Bjeluha, 1400 m, 24. 9. 1966, iïom Apodemus flavicollis, leg. B. Petrov id" i9 Kučište, near Peč ( = Rugovo, Kučište), 1350 m, from Glis glis, August 1939, E. Martino, presented by Zool. Mus. Hamburg, ex Wagner coll. (Brit. Mus. 1961. 677) (Hopkins & Rothschild, 1971) l9 Istok, 600 m, 6. 6. 1977, from Apodemus mystacinus epimelas, leg. B. Petrov Makedonija:
lCf Guri Vilpnis, Korab (=Korab: Guri Velpnis), [41° 45' N, 20° 40' E], from
Sylvaemus silvaticus stankovici (—Apodemus sylvaticus), July 1935, leg. E. Martino (Hopkins & Rothschild, 1971) 2(5 49 Sar Planina: Lukovo Pole, 1750 m, 7. 6. 1989, from Dtyomys nitedula, leg. B. Krvštufek
1(5 29 Bistra, 25. 7. 1933, from Sciurus vulgaris, leg. V Martino (»Ctenopsyllus sciuro-
bius«) (Wagner, 1934,1936,1939) id" Ničpur, Macedonia (=Ničpurska Planina: Ničpur), 13. 7. 1935, from Apodemus mystacinus epimelas, leg. V Martino (Hopkins & Rothschild, 1971) Wagner (1934) described this species on the basis of 3 specimens collected from S. vulgaris. According to present knowledge L. sciurobia is a separate species, which was not clear at the time of description. However it is not yet certain which is its major host. Apart from S. vulgaris it has been collected on the rodent species: D. nitedula, G. glis, A. flavicollis, A. sylvaticus, and/1, mystacinus. It occurs on small rodents nesting on the ground (Apodemus), also on those in tree hollows and nestboxes (Glis, Dtyomys), and in nests made of branches in tree canopies (Sciurus). According to Rosicky (1957) there are two different ecological groups of fleas, i.e. fleas of S. vulgaris and dormice, and fleas of small terrestrial mammals. This is interesting, because usually a single flea species belongs only to one ecological group. At the moment there are not enough data to clearly support this conclusion.
CERATOPHYLLIDAE
19.	Dasypsyllus (Dasypsyllus) gallinulae gallinulae (Dale, 1878) Slovenij a:
id" Snežnik: Sviščaki, 1250 m, 6. 10. 1994, from 1 nest of Glis glis, leg. T. Trilar
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20.	Myoxopsylla (Myoxopsylla) laverani laverani (Rothschild, 1911) Hrvatska:
i9 Krk Island: Veli vrh, 350 m, 5. 4. 1975, from Apodemus sylvaticus, leg. N. Tvrtkovič i9 Pag Island: Metajna, 20 m, 23. 7. 1988, from Eliomys quercinus, leg. B. Krystufek 7d 69 Pag Island: Pag, environs, 23. 7. 1977, from Eliomys quercinus, leg. N. Tvrtkovič 3d1 79 Velebit: Tulove grede, 890 m, 3. 8. 1975, from Eliomys quercinus, leg. N. Tvrtkovič
Id Ražanac, Vrankoviči, 40 m, 6. 7. 1975, from Eliomys quercinus, leg. N. Tvrtkovič l9 Hvar Island: Brusje, 340 m, 4. 10. 1975, from Eliomys quercinus, leg. N. Tvrtkovič 7 ex. Split, from 5 ex. Eliomyspallidus (=Eliomys quercinus), leg. J. Wagner (»Myoxopsylla laverani«) (Wagner, 1939)
Myoxopsylla I. laverani was found only within the range of E. quercinus, which is most probably the major host of this flea. Myoxopsylla l. laverani was not found in this area on G. glis, and therefore is not its major host, although it is found in other geographical areas on G. glis.
21.	Tcusopsylla octodecimdentcita octodecimdentata (Kolenati, 1863) Slovenija:
l9 Turjak, 500 m, 4. 12. 1977 from Martesfoina, leg. A. Šmuc id Poljanska dolina, 17. 9. 1968, from Sciurus vulgaris, leg. S. Brelih & F. Leben l9 Šenčur, Voglje, 370 m, 15. 8. 1986, from Sciurus vulgaris, leg. A. Kajzer l9 Ljubljana, environs, 300 m, 15. 12.1969, from Sciurus vulgaris, leg. S. Brelih 2d 49 Gameljne, 320 m, winter 1954, from Sciurus vulgaris, leg. J. Carnelutti
(»Tarsopsylla octodecimdentata«) (Rosicky & Carnelutti, 1959) 2d 29 Zgornje Jezersko, 1000 m, 12. 12. 1965, from Maries martes, leg. S. Brelih & A. Smuc
id l9 Litija, 300 m, 1. 12. 1971, from Martes martes, leg. F. Leben Hrvatska:
29 Dubrovnik, 1. 12. 1979, from Martes foina, leg. A. Lesinger Bosna i Hercegovina:
39 Igman, from Sciurus vulgaris, leg. V Martino (»Tarsopsylla octodecimdentatus«)
(Wagner, 1928-29, 1939) Jugoslavija: Srbija:
id 49 Tara Planina: Predov ICrst, 1100 m, 29. 10. 1968, from Sciurus vulgaris, leg.
B. Petrov Makedonija:
Bistra, from Sciurus vulgaris, leg. J. Wagner (»Tarsopsylla octodecimdentatus«) (Wagner, 1939)
The major host of T. o. octodecimdentata is S. vulgaris. It is very often found on M. martes and M. foina, because S. vulgaris is their frequent prey and because M. martes often choose the nests of S. vulgaris as a resting place.
Tarsopsylla o. octodecimdentata was found outside of Slovenia only at two sites, because from here very few specimens of S. vulgaris was examined.
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Fig. 9: Distribution of Spermophilus citellus (Linnaeus) (adapted from Petrov, 1992), and survey sites of Ctenophthalmus orientalis (Wagner)
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Fig. 10: Distribution of Eliomys quercinus (Linnaeus) (adapted from Petrov, 1992 and Tvrtkovic et al., 1995), and survey sites of Tarsopsylla octodecimdentata octodecimden-tata (Kolenati), Myoxopsylla laverani laverani (Rothschild), Echidnophaga murina (Tiraboschi), and Leptopsylla sciurobia (Wagner)
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22. Citellophilus simplex simplex (Wagner, 1902) Jugoslavia: Srbija: Vojvodina:
id 49 Deliblatska Pescara: Samos, 100 m, 14. 5. 1989, from Spemiophilus citellus, leg. B. Krystufek
1id 59 Deliblatska pescara (correct Deliblatska Pescara: Deliblato), 7. 5. 1947, from Citellus citellus (=Sperinophilus citellus), leg. A. Ruzic (Rosicky & Todorovic, 1964) 39 Deliblato (=Deliblatska Pescara: Deliblato), 100 m, May 1945, from Citellus citellus, (=Spermophilus citellus), leg. A. Ruzic (Rosicky & Todorovic, 1964) 12d 149 Deliblatska Pescara: Deliblato, 100 m, 5. 4. 1949, from Spermophilus citellus,
leg. A. Ruzic 5d 59 idem, 20. 4. 1949, leg. B. Petrov id l9 idem, 21. 4. 1949
id Deliblatska Pescara: Susara, 170 m, 13. 5. 1989, from Spermophilus citellus, leg. B. Krystufek
lCf 89 Banat: Banatska Palanka, 85 m, 28. 6. 1988, from Spermophilus citellus, leg. B.
Krystufek id 29 idem, 29. 6. 1988 id 69 idem, 30. 6. 1988
4d ll9 idem, 15. 4. 1989, leg. B. Krystufek & T. Trilar Makedonija:
l9 Dojransko jezero, Acikot, 150 m, 31. 5. 1988, from Spermophilus citellus, leg. B.
Krystufek id idem, 8. 5. 1989 id l9 idem, 9. 5. 1989
Two species of the genus Citellophilus are distributed in Europe, C. simplex and C. martinoi. Both species are easily distinguishable, but infraspecific distinctions are very complicated. Almost all taxonomic characters are highly variable, especially the movable process of the clasper. There are some similarities between geographically neighboring populations, but there are also large differences within the same population. Cyprich (1989) discussed this subject in a revision of a large amount of material, and our statements are based on his findings.
Citellophilus simplex is described from the Ukraine and is distributed from Turkey and southern Russia, across Moldavia, Romania, Bulgaria, as far as Yugoslavia (southeastern Vojvodina), Hungary, and Slovakia. In eastern Slovakia both subspecies occur: the nominate subspecies is limited to the extreme eastern part of Slovakia, while C. s. rosiclcyi Cyprich occurs west from there in a limited area (Fig. 20). We found the nominate subspecies in many sites in southeastern Vojvodina (Deliblatska Pescara, Banatska Palanka) and in Macedonia (Dojran), which is the first finding in Macedonia.
Citellophilus simplex has two major hosts: S. suslicus in eastern Europe and S. citellus in
southern and southeastern Europe.
We further discuss this topic in the Discussion section.
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Fig. 11: Citellophilus simplex simplex (Wagner): process of clasper of male: a. Banatska Palanka, Banat, Vojvodina; b. Jazak, Fruska Gora, Srem, Vojvodina
Fig. 12: Citellophilus martinc martino. (Wagner & Ioff): process of clasper of male: a. Vlasina, southeastern Serbia; b. Begovo Pole, Mt. Jakupica, Macedonia
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Fig. 13: Citellophilus simplex simplex (Wagner): sternum VII of female and outline of sternum VII of female: a. Banatska Palanka, Banat, Vojvodina; b. Ačikot, Dojran, Macedonia
Fig. 14: Citellophilus martinoi martinoi (Wagner & Ioff) sternum VII of female and outline of sternum VII of female: a. Jazak, Fruska Gora, Srem, Vojvodina; b. Begovo Pole, Mt. Jakupica, Macedonia
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Fig. 15: Spermatheca of female of: a, b. Citellophilus simplex simplex (Wagner) (a. Samoš, Deliblatska Peščara, Vojvodina; b. Ačikot, Dojran, Macedonia); and c, d. C telloph (us martinoi martinoi (Wagner & Ioff) (c. Jazak, Fruška Gora, Srem, Vojvodina; d. Begovo Pole, Mt. Jakupica, Macedonia)
23. Citelloph us martini martinoi (Wagner & Ioff, 1926) Jugoslavija: Srbija: Vojvodina:
I rig, Srem ( = Srem: Fruška Gora: I rig), from Ci telli is citellus (-Spermophilus citellus),
24. 9. 1924, leg. V Martino (»Ceratophyllus martinoi«) (Wagner, 1928-29) 2d 69 Srem: Fruška Gora: Jazak, 180 m, 12. 4. 1989, from Spermophilus citellus, leg.
B. Krystufek & T. Trilar 29 Bačka: Titel, Lok, 80 m, 14. 5. 1989, from Spermophilus citellus, leg. B. Krystufek Melenci, Bačka (correct Banat: Melenci), from Citellus citellus (=Spermophilus citellus), leg. V. Martino (»Ceratophyllus martino <) (Wagner, 1928-29) ltf 29 Banat: Zrenjanin, Orlovat, reka Tamiš, 81 m, 13. 4. 1989, from Spermophilus
citellus, leg. B. Krystufek & T. Trilar Jugoslavija: Srbija:
l9 Vlasinska Tresava, 1200 m, 18. 5. 1948, from Spermoph lus citellus, leg. B. Petrov 169 Vlasina, 1250 m, August 1947, from Citellus citellus (=Spermophilus citellus), leg. A. Ružič (Rosicky & Todorovič, 1964) 15CJ 319 ibidem, 18. 8. 1947, from Talpa europaea, leg. A. Ružič (Rosicky & Todorovič, 1964)
10Cf 329 ibidem, 25. 7. 1977, from Spermophilus citellus, leg. B. Petrov IOCS 169 Stara Planina: Ponor, 17. 6. 1947, from Citellus citellus (—Spermophilus citellus), leg. A. Ružič (Rosicky & Todorovič, 1964)
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Citellophilus martinoi, like C. simplex, is a highly variable species. There are morphological similarities especially between geographically neighboring populations, but at the same time there is marked high intrapopulation variation. Citellophilus martinoi is described from Irig (Frušlca Gora, Srem) and Melenci (Banat; Bačka is incorrectly cited in the literature), both loca typica are in Vojvodina (Yugoslavia). The nominate subspecies occurs also in Bulgaria, Romania, east Serbia, south Hungary, and in the central part of Slovakia (Fig. 20). According to Cyprich (1989) the subspecies C. m. rotundus Rosicky is distributed in the Czech Republic, Moravia, part of Slovakia, Hungary and in eastern Austria. Transitional forms between the subspecies are frequent.
The major host of C. martinoi is S. citellus. In eastern Serbia it was collected on T. europaea. As other species of the genus Cittelophilus, it lives in the fur of the host.
Citellophilus martinoi ssp. Makedonija:
3C? 159 Jakupica: Gorno Begovo, Begovo Pole, 1950-2000 m, 11. 5. 1989, from
Spermophilus citellus, leg. B. Krystufek 29 idem, 27. 6. 1989 2(5 l9 idem, 28. 6. 1989
1(5 59 Jakupica: Solunsko Pole, 2100 m, 28. 6. 1989, from Spermophilus citellus, leg. B. Krystufek
Krystufek collected this material in 1989 in an isolated mountain population of S. citellus (S. c. karamani) at the Mt. Jakupica in Macedonia. This was the first finding of C. martinoi in Macedonia. The collected 6 males and 23 females were not sufficient for a final determination at the subspecific level. There is also substantial intrapopulation variation. The movable process of the clasper is quite narrow, otherwise there are no relevant differences. Further studies are needed to determine whether they should be considered to represent the nominate subspecies or not.
Citellophilus sp.
Makedonija
Kožuf, from Citellus citellus {—Spermophilus citellus), leg. J. Wagner (»Citellophilus martinoi.« - with the note that it could be another species) (Wagner, 1939) Wagner probably collected the material somewhere at the foot of the Mt. Kožuf, while S. citellus does not live at the mountain.
24. Ceratophyllus (Monopsyllus) sciurorum sciurorum (Schrank, 1803) Sloven i j a:
5(5 29 Komen, Kobjeglava, 324 m, 30. 7. 1985, from Glisglis, leg. B. Petrov 2(5 29 Trnovski gozd: Čaven, 1240 m, 3. 10. 1968, from Glisglis, leg. S. Brelih & R. Jelinčič
l9 ibidem, 10. 7. 1969, from Apodemus sylvaticus, leg. S. Brelih & B. Petrov 32C? 319 Hotedršica (=Hotedršica, Novi svet), 700 m, 26. 8. 1956, from Glisglis, leg. R. & N. Jelinčič (»Monopsyllus sciurorum«) (Rosicky & Carnelutti, 1959)
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Fig. 16: Distribution of Spermophilus citellus (Linnaeus) (adapted from Petrov, 1992), and survey sites of Citellophilus martinoi martinoi (Wagner & Ioff) (included Citellophilus martinoi ssp. from the Mt. Jakupica in Macedonia), and Citellophilus simplex simplex (Wagner)
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3d 39 Hotedršica, Novi Svet, 650 m, 4. 8. 1967, from Glis glis, leg. S. Brelih & R. Jelinčič
4d 29 Hotedršica, Ravnik, 650 m, 5. 8. 1967, from Glis glis, leg. S. Brelih & R. Jelinčič ld Snežnik, 1600 m, 25. 9. 1968, from Glis glis, leg. F. Leben
3d 179 Snežnik: Sviščaki, 1250 m, 2. 9. 1992, from 1 nest of Muscardinus avellanarius, leg. T. Trilar
2d IV ibidem, 1. 6. 1993, from 1 nest of Muscardinus avellanarius
2d 29 Borovnica, Pako, 400 m, 15. 9. 1968, from Glis glis, leg. F. Leben
l9 Turjak, 500 m, 4. 12. 1977, from Martesfoina, leg. A. Šmuc
92d 1849 Goteniška gora: Draga, 840 m, 9. 9. 1992, from 3 nests of Glis glis, leg. T.
Trilar iTrilar, 1995,1997) 23ld 2589 idem, 29. 9. 1998, from 3 nests of Glis glis, leg. K. Koselj l9 idem, from 1 nest of Muscardinus avellanarius
l9 idem, 29. 9. 1998, from 1 nest of Muscardinus avellanarius, leg. K. Koselj 3d 29 idem, 6. 11. 1996, from 1 nest of Glis glis, leg. T. Trilar & M. Perušek id 29 Kočevje, environs, 15.10. 1958, from Glis glis, leg. S. Brelih 3d 49 Bohinj, Ukane, 560 m, 28. 7. 1957, from Glis glis, leg. S. Brelih (»Monopsyllus
sciurorum«) (Rosicky & Carnelutti, 1959) 2d 39 Trenta: Julijana, 720 m, 13. 9. 1968, from Glis glis, leg. S. Brelih & J. Dovič l9 Rateče, Zelenci, 834 m, 14. 7. 1974, from Apodemus flavicollis, leg. S. Brelih & J. Gregori
id Cerkno, 400 m, 6. 2. 1969, from Sciurus vulgaris, leg. F. Leben 7d 79 Jelovica: Goška ravan, 970 m, 11. 9. 1973, from Glis glis, leg. B. Kryštufek 170d 2979 Rudno, 680 m, 3. 11. 1999, from 1 nest of Glis glis, leg. T. Trilar 105d 2049 Rudno, Hujska, 710 m, 3. 11. 1999, from 1 nest of Glis glis, leg. T. Trilar 29 Kranj, Sveti Jošt, 800 m, 15. 5. 1983, from Glis glis, leg. J. Perovič Tržič, series from Sciurus vulgaris, leg. S. Brelih & D. Tovornik (»Monopsyllus sciurorum«) (Rosicky & Carnelutti, 1959) 30d 10l9 Podljubelj, Čižovnik, 950 m, 5.11.1999, from 1 nest of Glis glis, leg. T. Trilar
l9 Ljubljana: Vrhovci, 300 m, 17. 1. 1972, from Sciurus vulgaris, leg. J. Dovič l9 Medvode, 320 m, 1. 3. 1956, from Vulpes vulpes, leg. J. Carnelutti (»Monopsyllus
sciurorum«) (Rosicky & Carnelutti, 1959) l9 Sorško polje, Žabnica, 370 m, 3. 11. 1999, from 1 nest of Glis glis, leg. T. Trilar 6d 169 Cerklje, Stefanja gora, 750 m, 28. 6. 1975, from nest of Parus major, leg. D. Sere
Kranj, 385 m, from Glis glis, leg. L. Kuščer (»Ceratophyllus sciurorum«) (Wagner, 1928-29)
id l9 Kranj, Milje, 420 m, 20. 3. 1992, from Sciurus vulgaris, leg. I. Zavrl 3d J 9 ibidem, 30. 9. 1973, from Glis glis, leg. B. ICrystufek l9 Šenčur, Voglje, 370 m, 25. 8. 1986, from Sciurus vulgaris, leg. A. Kajzer 4d 59 Lom, Grahovše, Gaberčev rovt, 1290 m, 5. 11. 1999, from 1 nest of Glis glis, leg. T. Trilar
4d 29 Črnuče, 320 m, from Glis glis, leg. J. Carnelutti (»Monopsyllus sciurorum«)
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(Rosicky & Carnelutti, 1959) l9 Ljubljana, 300 m, 15. 5. 1983, horn Homo sapiens, leg. I. Sivec 2 a Gameljne, 320 m, winter 1954, from Sciurus vulgaris, leg. J. Carnelutti
(»Monopsyllus sciurorum«) (Rosicky & Carnelutti, 1959) Kamnik, environs, big series from Glis glis, leg. J. Wagner (»Ceratophyllus sciurorum«) (Wagner, 1928-29)
idem, from Sciurus vulgaris {»Ceratophyllus sciurorum«) (Wagner, 1928-29) ICS Kamniška Bistrica, 600 m, from Apodemus flavicollis, 26. 6. 1957, leg. Czechoslovak Slovenian expedition (»Monopsyllus sciuronun«) (Rosicky & Carnelutti, 1959)
Jezersko ( = Zgornje Jezersko), from Eliomys quercinus (correct Diyomys nitedula), leg.
J. Wagner (»Ceratophyllus sciurorum«) (Wagner, 1928-29) idem, from Evotomys glareolus (=Clethrionomys glareolus) (»Ceratophyllus sciurorum«) (Wagner, 1928-29)
ICS Zg ornje Jezersko, 1000 m, 12. 12. 1965, from Martes martes, leg. A. Smuc 4C1 99 Litija, 300 m, 1. 12. 1971, from Maries martes, leg. F. Leben 8CS ll9 Mirna na Dolenjskem, 300 m, 28. 4. 1969, from Maries martes, leg. S. Brelih l9 Kostanjevica, Šentjernej, 159 m, 11. 2. 1976, from Felis silvestris, leg. A. Šmuc YllCS 1959 Slovenj Gradec, 460 m, 20. 9. 1996, from 1 nest of Glis glis, leg. T. Novak ICS Pohorje: Lobnica, 1000 m, 18. 7. 1969, from Clethrionomys glareolus, leg. S. Brelih & B. Petrov
l9 Kozje, Lesično, Gruska jama, 16. 10. 1978, found in the pitfall, leg. T. Novak & I. Sivec
l9 Polj čane, Studenice, Jama v kamnolomu, 260 m, 16. 10. 1978, found in the pitfall,
leg. T. Novak & I. Sivec i9 Dvorjane, Vurberg, Grmada, 360 m, 8. 5. 1994, from Micromys minutus, leg. F. Janžekovič
1 CS Srednja Bistrica, 170 m, 19. 6. 1957, from Clethrionomys glareolus, leg. Czechoslovak Slovenian expedition (»Monopsyllus sciuronun«) (Rosicky & Carnelutti, 1959) Hrvatska:
Babja Gora, Slavonija (=Slavonija: Babja Gora), from Sciurus vulgaris, leg. V Martino
(»Ceratophyllus sciurorum«) (Wagner, 1928-29) ICS Velebit: Mirevo, 1400 m, 15. 8. 1976, from Diyomys nitedula, leg. N. Tvrtkovič 2CS idem, 7. 8. 1977
39 Velebit: Zavižan, 1594 m, 24. 6. 1971, from Glis glis, leg. G. Džukič 4CS 79 Velebit: Zavižan, Modrica dolac, 1460 m, 10. 7. 1984, from Diyomys nitedula, leg. A. Vukušič
l9 Gorski Kotar: Risnjak, 1400 m, 9. 8. 1967, from Diyomys nitedula, leg. S. Brelih & B. Petrov
2CS l9 Velebit: Baške Oštarije, 900 m, 11. 8. 1968, from Eliomys quercinus, leg. S. Brelih
ICS ibidem, 925 m, 26. 6. 1981, from Apodemus flavicollis, leg. N. Tvrtkovič l9 Velebit: Papratnjak, 920 m, 30. 7. 1976, from Diyomys nitedula, leg. N. Tvrtkovič
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l9 Velebit: Bunjevac, 1200 m, 15. 6. 1986, many specimens on the soil, leg. B. Jalžič 2d ibidem, 7. 8. 1977, from Dryomys nitedula
2(5 Velebit: Predzid, 800 m, 2. 8. 1975, from Glisglis, leg. N. Tvrtkovič & B. Krystufek id 29 ibidem, 740 m, 2. 8. 1975, from Eliomys quercinus, leg. N. Tvrtkovič & B. Krystufek
id Vukomeričke gorice: Prkovec, 200 m, 10. 8.1974, from Glisglis, leg. N. Tvrtkovič ld Turopolje: Peščenica, near the river Odra, 95 m, 7. 4. 1974, from Apodemus
flavicollis, leg. N. Tvrtkovič 3C1 39 Zagreb, Dugo Selo, 100 m, 5. 4.1959, from Martes martes, leg. K. Igalffy 39 Brač Island: Dol, 600 m, 4. 9. 1975, from Glisglis, leg. N. Tvrtkovič 9d 39 idem, from Apodemus sylvaticus
9(5 89 Biokovo: Sveti Jura, 1650 m, 31. 7. 1970, from Glisglis, leg. B. Petrov & G.
Džukič Bosna i Hercegovina:
Sjetlino, Bosna (=Sjetlino), from Glisglis, leg. Zjuzin (»Ceratophyllus sciurorum«) (Wagner, 1928-29)
id" 29 Bjelašnica (Gacko), 1600 m, 4. 8. 1970, from Glisglis, leg. B. Petrov
Igman, from Glis glis, leg. V Martino (»Ceratophyllus sciurorum«) (Wagner, 192829) idem, from Sciurus vulgaris (»Ceratophyllus sciurorum«) (Wagner, 1928-29) 2d Zelengora: Orlovačko jezero, 1500 m, 9. 9. 1984, from Diyomys nitedula, leg. B. Krystufek
id" Gazivoda, 26. 10. 1988, from Glis glis, leg. B. Krystufek 2(5 49 Šator planina: Šatorsko jezero, 1490 m, 20. 9. 1988, from Glisglis, leg. B. Krystufek
5(5 20 Šator planina: Babina greda, 1620 m, 5. 9. 1984, from Glisglis, leg. B. Krystufek id Svinjača, Blidinje jezero, 1170 m, 20. 6. 1971, from Microtus aivalis, leg. G. Džukič 2d Velež, Rujište, 1050 m, 18. 5. 1971, from Glisglis, leg. B. Petrov Jugoslavija: Črna Gora:
39 Hercegnovi, 50 m, 15. 6.1990, from Diyomys nitedula, leg. B. Krystufek Cetinje, Črna Gora ( = Cetinje), from Glis glis, leg. V Martino (»Ceratophyllus sciurorum«) (Wagner, 1928-29) l9 Orjen, 1100 m, 13. 10. 1990, from Sciurus vulgaris, leg. B. Krystufek l9 Bjelasica: Zekova Glava, 2050 m, 19. 9. 1981, from Dinaromys bogdanovi, leg. B.
Petrov & M. Milenkovič (Brelih, 1986) Jugoslavija: Srbija: Vojvodina:
l9 Srem: Erdevik, 119 m, 29.10. 1963, from Apodemus flavicollis, leg.D. & N. Heneberg
ld Srem: Fruška Gora: Zmajevac, 450 m, 28. 10. 1963, from Apodemus flavicollis, leg. D. & N. Heneberg
ld Deliblatska Peščara: Kremenjak, 100 m, 27. 2. 1987, from Sciurus vulgaris, leg. M.
Milenkovič Jugoslavija: Srbija:
4d l9 Tara Planina: Kremiči, 950 m, 20. 6. 1949, from Glis glis, leg. B. Petrov 2d 29 Tara Planina: Predov Krst, 1100 m, 29. 10. 1968, from Sciurus vulgaris, leg. B.
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Petrov
2d l9 Golija: Biser Voda, 1200 m, 19. 6. 1970, from Glis glis, leg. B. Petrov Arandelovac, from Glis glis, leg. V Martino (»Ceratophyllus sciurorum«) (Wagner, 1928-29)
id Niš: Sičevo, 13. 5. 1970, fr om Apodemus sylvaticus, leg. B. Petrov l9 Veliki Jastrebae, Ravnište, 600 m, 14. 6. 1979, from Apodemus flavicollis, leg. B. Petrov
5d 59 Suva Planina, 7. 6. 1947, from Rattus rattus, leg. B. Petrov {»Monopsyllus sciurorum«) (Rosicky & Todorovič, 1964) 3CJ idem, 9. 6. 1947, from Sciurus vulgaris {»Monopsyllus sciurorum«) (Rosicky & Todorovič, 1964)
Donji Milanovac, from Glis glis, leg. V Martino {»Ceratophyllus sciurorum«) (Wagner, 1928-29)
i9 Besna Kobila, ~ 1600 m, August 1947, from Mus musculus, leg. B. Petrov
{»Monopsyllus sciurorum«) (Rosicky & Todorovič, 1964) 2d 39 Vlasina, 1250 m, August 1947, from Sciums vulgaris, leg. B. Petrov
{»Monopsyllus sciurorum«) (Rosicky & Todorovič, 1964) id l9 idem, August 1947, from Glis glis {»Monopsyllus sciurorum«) (Rosicky & Todorovič, 1964)
2d Suva Planina: Tri lokve, 1400 m, 14. 8. 1981, from Glis glis, leg. B. Petrov 2d idem, from Rattus rattus
l9 Vidlič: Basara: Planinica, 1200 m, 17. 8. 1981, from Glis glis, leg. B. Petrov lld 109 Stara Planina: Dojkinci, ~ 900 m, 25. 6. 1947, from Glis glis, leg. B. Petrov
{»Monopsyllus sciurorum«) (Rosicky & Todorovič, 1964) l9 Stara Planina: Dojkinci, Bašta, 13.-17. 10. 1947, from Apodemus sylvaticus, leg. B.
Petrov {»Monopsyllus sciurorum«) (Rosicky & Todorovič, 1964) ld 29 Stara Planina, October 1947, from Glis glis, leg. B. Petrov {»Monopsyllus sciurorum«) (Rosicky & Todorovič, 1964) 2d 59 idem, 27. 6. 1947 (»Monopsyllus sciuronun«) (Rosicky & Todorovič, 1964) 3d 49 idem, September 1947, from Sciurus vulgaris (»Monopsyllus sciurorum«)
(Rosicky & Todorovič, 1964) 8d idem, 21. 6. 1947 {»Monopsyllus sciurorum«) (Rosicky & Todorovič, 1964) l9 Stara Planina: Kopren, ~ 1850 m, 29. 6. 1947, from Apodemus sylvaticus, leg. B.
Petrov {»Monopsyllus sciurorum«) (Rosicky & Todorovič, 1964) Jugoslavija: Srbija: Kosovo:
9d 29 Žljeb: Kula, 1600 m, 24. 6. 1980, from Glis glis, leg. B. Petrov id idem, 1750 m, 14. 6. 1977, from Diyomys nitedula Makedonija:
Bistra, 25. 7. 1933, series from Sciums vulgaris, leg. V Martino (»Monopsyllus sciurorum«) (Wagner, 1934) 29 Kičevo, Dolenci, 620 m, 15. 9. 1989, from Glis glis, leg. B. Krystufek l9 Šar Planina: Jelak, 1900 m, 21. 7. 1978, from Apodemus flavicollis, leg. G. Džukič 3d Prilep, 700 m, 21. 4. 1969, from Diyomys nitedula, leg. B. Petrov
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Fig. 17: Distribution of Ceratophyllus sciurorum sciuromm (Schrank) in the Western and Central Balkans
Ceratophyllus s. sciuromm is the most common flea species of our red squirrels and dormice, especially G. glis. It often passes to other hosts, which come in contact with squirrels and dormice as predators or during nesting. It is a widespread sylvatic flea in the Western and Central Balkans found on the following hosts: S. vulgaris, G. glis, D. nitedula, E. quercinus, M. avellanarius, M. martes, M. foina, V vulpes, F. silvestris, A. flavicollis, A. sylvaticus, M. musculus s. lat., M. minutus, R. rattus, C. glareolus, M. aivalis, D. bogdanovi, in the nest of P. major, and also on humans, on floors, the soil, and even in carstic caves in traps with beetle bait.
25. Ceratophyllus (Monopsyllus) sp.
SI oven ij a:
l9 Snežnik: Sviščaki, 1250 m, 22. 7. 1997, from 1 nest of Glis glis, leg. T. Trilar 2Ci ibidem, 3. 9. 1999
2Cf ibidem, 30. 10. 1999, from 2 nests of Glis glis
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During the determination of a great amount of material from the nests of G. glis from Sviščaki, Mt. Snežnik (Slovenia) (Tab. 1) we found 4 males and 1 female, which are clearly distinguishable from the all known species from the genus Ceratophyllus. The males differ in the shape of sternum VIII and females in the shape of sternum VII and in the shape of spermatheca. These most likely represent an undescribed species.
26.	Ceratophyllus (Ceratophyllus) gallinae (Schrank, 1803) Slovenija:
l9 Snežnik: Sviščaki, 1250 m, 24. 10. 1997, from 1 nest of Glis glis, leg. T. Trilar l9 ibidem, 16. 7. 1999, from 1 nest of Glis glis
1C? Podljubelj, Čižovnik, 950 m, 5. 11. 1999, from 1 nest of Glis glis, leg. T. Trilar
Ceratophyllus gallinae ecologically belongs to the group of fleas of small passeriform birds which nest in tree canopies, holes, nest boxes, and bushes (Rosicky, 1950, 1957; Jurik, 1975,1976,1978). Glis glis is an occasional host (Traub et al., 1983).
27.	Ceratophyllus (Ceratophyllus) hirundinis (Curtis, 1826) Slovenija:
3C? 39 Snežnik: Sviščaki, 1250 m, 22. 8. 1990, from 1 nest of Glis glis, leg. T. Trilar (Trilar, 1995, 1997)
28.	Ceratophyllus (Ceratophyllus) rusticus Wagner, 1903 S1 oven ij a:
l9 Snežnik: Sviščaki, 1250 m, 22. 8. 1990, from 1 nest of Glis glis, leg. T. Trilar (Trilar, 1995,1997)
The presence of C. hirundinis and C. rusticus in G. glis nests was very surprising (Trilar, 1995, 1997). Both belong to the group of swallow fleas (Rosicky, 1950, 1957; Jurik, 1975, 1976, 1978), and G. glis is an accidental host (Traub et al, 1983). Which intermediate host mediates transfer from swallows to dormice is unknown, because swallows nest at great distances from rodent dens.
Discussion
We have studied fleas (Siphonaptera) from the territory of the Western and Central Balkans (former Yugoslavia) (Fig. 1) whose hosts are rodents from the family of squirrels (Sciuridae) and dormice (Gliridae). Three species of squirrels live in this area: the red squirrel (S. vulgaris), the European souslik (S. citellus), and the Alpine marmot (M. marmota), and four species of dormice: the fat dormouse (G. glis), the common dormouse (M. avellanarius), the garden dormouse (E. quercinus), and the forest dormouse (D. nitedula). Tables 2 trough 7 provide results of our survey of fleas found on S. vulgaris, S. citellus, G. glis, M. avellanarius, E. quercinus, and D. nitedula in the Western and Central Balkans.
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Date	Nests No.		
6. 3. 1990	6	226	311
2. 6. 1990	1	1	8
22. 8. 1990	4	190	221
30. 6. 1992	8	649	752
2. 9. 1992	12	556	614
17.2. 1993	11	249	432
27. 9. 1993	3	21	34
22. 6. 1994	7	4	10
6. 10. 1994	8	417	562
4. 7. 1995	10	150	294
1. 8. 1996	7	88	122
19. 9. 1996	3	469	659
22. 7. 1997	3	81	86
24. 10. 1997	5	176	152
9. 10. 1998	6	129	122
28. 5. 1999	2	0	6
16. 7. 1999	9	64	107
3. 9. 1999	8	110	166
30. 10. 1999	16	569	1039
Total	83	3781	5166
Tab. 1: Survey of Ceratophyllus sciurorum sciuronun in the nests of Glis glis from Sviščaki, Mt. Snežnik (1250 m, leg. T. Trilar) data from 1990 to 1993 according to Trilar (1995, 1997).
		S		HR		BH		MTG		VOJ		SR	B	KOS		MK	
		No	L	No	L	No	L	No	L	No	L	No	L	No	L	No	L
L sciurobia	3 ?															1 2	1
T. octodecimdentata octodecimdentata	6 ?	3 5	4			3	1					1 4	1			+	V
C. sciurorum sciurorum	<J $	3 3	4 2*	+	r	+	V	1	1	1	1	7 5	4			+	V
Tab. 2: Survey of fleas on Sciurus vulgaris Linnaeus in the Western and Central Balkans:
No - Number of specimens L - Number of UTM squares with survey sites
+ - Species present, but without a citation of the number of specimens - Number of UTM squares, without a citation of the number of specimens
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		S		HR		BH		MTG		VOJ		SR	B	KOS		MK	
		No	L	No	L	No	L	No	L	No	L	No	L	No	L	No	L
N. setosa spinea	<J 9									5 11	4						
C. agyrtes graecus	6 9															1 1	2
C. orientalis orientalis	S ?									15 21	8 1*	10 19	3				
C. orientalis jakupicae	6 9															24 33	1 1*
Citellophilus sp.	<3 9															+	V
C. simplex simplex	6 9									38 59	4					2 2	4
C. martinoi martinoi	6 9									3 10	3 2*	22 65	3			6 23	
C. martinoi ssp.	6 9															6 23	1
Tab. 3: Survey of fleas on Spermophilus citellus (Linnaeus) in the Western and Central Balkans:
No - Number of specimens L - Number of UTM squares with survey sites
+ - Species present, but without a citation of the number of specimens - Number of UTM squares, without a citation of the number of specimens
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		SI		HR		BH		MTG		VOJ		SRB		KOS		MK	
		No	L	No	L	No	L	No	L	No	L	No	L	No	L	No	L
Pulex irritans	6 ?	1	1														
H. orientalis orientalis	c? ?					1	1										
C. monticola	S 9											1 1	2				
C. agyrtes wagnerianus	c? ?	3	2														
C. agyrtes ohridanusodinarus	?			4 6	1												
C. nifetodes brelihi	?	3 4	1														
C. nifetodes tvrtkovici	6 ?			1	1												
C. nifetodes nifetodes	6 $					2	1	1	1								
C. congener congener	6 ?	1	1														
L segnis	6 ?			1 2	1												
L sciurobia	<J ?													1 1	1		
C. sciurorum sciurorum	c? ?	4662 6476	21 2*	12 14	4	11 8	4 2*	+	1*			34 25	7 2*	g 2	1	2	1
C. hirundinis	$	3 3	1														
C. rusticus	a ?	1	1														
Tab. 4: Survey of fleas on Glis glis (Linnaeus) in the Western and Central Balkans: No - Number of specimens L - Number of UTM squares with survey sites
+ - Species present, but without a citation of the number of specimens - Number of UTM squares, without a citation of the number of specimens
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		S		HR		BH		MTG		VOJ		SR	B	KOS		MK	
		No	L	No	L	No	L	No	L	No	L	No	L	No	L	No	L
C. sciurorum sciurorum	6 $	5 20	2														
Tab. 5: Survey of fleas on Muscardimts avellananus (Linnaeus) in the Western and
Central Balkans:
No - Number of specimens
L - Number of UTM squares with survey sites
+ - Species present, but without a citation of the number of specimens - Number of UTM squares, without a citation of the number of specimens
		SI		HR		BH		MTG		VOJ		SRB		KOS		MK	
		No	L	No	L	No	L	No	L	No	L	No	L	No	L	No	L
E. murina	6 ?			2	1												
M. laverani laverani	3 ?			11 15	5 1*												
C. sciurorum sciurorum	6 $			3 3	2												
Tab. 6: Survey of fleas on Eliomys quercinus (Linnaeus) in the Western and Central Balkans:
No - Number of specimens: L - Number of UTM squares with survey sites
+ - Species present, but without a citation of the number of specimens Number of UTM squares, without a citation of the number of specimens
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		SI		HR		Bh		MTG		VOJ		SR	B	KOS		MK	
		No	L	No	L	No	L	No	L	No	L	No	L	No	L	No	L
C. agyrtes graecus	<2 ?															1	1
C. congener troilus	6 ?															1	1
L. sciurobia	C? 9					1 1	2	1	1							2 4	1
C. sciurorum sciurorum	6 ?	+	1*	9 9	4	2	1	3	1					1	1	3	1
Tab. 7: Survey of fleas on Diyomys nitedula (Pallas) in the Western and Central Balkans:
No - Number of specimens L - Number of UTM squares with survey sites
+ - Species present, but without a citation of the number of specimens Number of UTM squares, without a citation of the number of specimens
M. marmota was introduced into the Alps (Julijske Alpe - Julian Alps), Kamnisko-Savinjske Alpe (Kamnik-Savinja Alps) and Karavanke (Karawanken) in northwestern Slovenia beginning in 1953. Indigenous populations of M. marmota were already extinct in this area at the end of the Pleistocene or the beginning of the Holocene (Krystufek, 1991). We did not examine any individuals or nests of this species so we do not have data on their ectoparasites.
Fleas in Europe based on hosts belong to thirteen ecological groups, where the systematic relationships are not important (Rosicky, 1957). The fleas from these hosts ecologically belong to the flea group of large ground rodents (Cricetus, Spermophilus) and the fleas of red squirrels and dormice but there are also others in tree canopies and tree hollow nesting birds and mammals.
The most common flea from the ecological group of red squirrels and dormice is C. s. sciurorum. Its major hosts are S. vulgaris and G. glis. All examined individuals and nests of both hosts were infested with C. s. sciurorum. This flea was also common on D. nitedula, but surprisingly rare on M. avellanarius and E. quercinus. It was often found on M. martes and once on M. foina, as these predators visit nests of squirrels and dormice. It was found on C. glareolus, D. bogdanovi, M. nivalis, M. minutus, M. muscu-lus s. lat., A. flavicollis, and A. sylvaticus, all of which are hosts of fleas which are belonging to the ecological group of small terrestrial mammals that nest on the ground. In Slovenia it has also been found in the nests of P. major. Some were also found on the floor of underground caves and buildings. We presume that fleas were
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transferred to such places by G. glis, a species known to use both caves (Polak, 1997) and buildings as shelter. Ceratophyllus s. sciwonun can also pass to humans and can also bite. It seems likely that in human habitations they also passed to R. rattus and M. musculus s. lat. In Slovenia, C. sciuronun was found also on two large predators, V vulpes and F. silvestris.
Tarsopsylla o. octodecimdentata and M. I laverani also occur on squirrels and dormouse. The major host of the first is S. vulgaris and are often passed to the predators M. martes and M. foina. This conclusion is based on our findings from Slovenia because only a few S. vulgaris from other areas of the Western and Central Balkans were examined. In general T. o. octodecimdentata is much rarer than C. s. sciurorum.
The major host of M. I. laverani is not known (Rosicky, 1957). In the Western and Central Balkans all the specimens were collected within the range of E. quercinus. Thus all the findings are from E. quercinus and a single female from A. sylvaticus. Although a great number of specimens from the Balkans were examined, M. I. laverani was never collected from G. glis which is usually cited as a major host. Our results suggest that E. quercinus is the major host of M. I. laverani. In Central Europe and the Balkans where G. glis and E. quercinus occur in the same habitats, it is possible that M. I. laverani passes from one host to another.
The first known host of L. sciurobia was S. vulgaris, which also gave it its name. It was also collected on G. glis and D. nitedula. Based on this data it belongs ecologically to the red squirrel and dormouse group of fleas. But it was also collected from A. sylvaticus and A. mystacinus and therefore could be regarded as belonging to the ecological group of fleas of small terrestrial mammals which also include the majority of species of Leptopsylla. The distribution of L. sciurobia includes the Central and Southern Balkans (Montenegro, Kosovo, Macedonia, Albania, and Greece). However the data are insufficient to determine their ecological affinity at present time.
From the ecological group of fleas of large ground rodents (Cricetus, Spermophilus) we will discuss only fleas from S. citellus, as the common hamster (C. cricetus) belongs to the family Cricetidae.
Spermophilus citellus is the major host of four flea species: N. setosa, Ct. orientalis, C. simplex, and C. martinoi. The first two species are typically found in nests and the latter two from fur. The distribution of C. simplex and C. martinoi is very complex.
Spermophillus citellus is endemic to Europe. The range is divided by the Carpatians into two main parts (Fig. 18). One includes the Pannonian basin and adjacent plains of the Czech Republic, Austria, Hungary, Slovakia, western Romania, and Yugoslavia. The other is to the south and east of the Carpatians in southern Romania, Bulgaria, Thrace, Moldavia, and Ukraine. Small peripheral isolated populations are present in Slovakia, Macedonia, and northern Greece (Krystufek, 1996; Mitchell-Jones et al., 1999), as well as recently extinct populations from southern Poland and eastern Germany. Eight subspecies were described, but authors disagrees as to infraspecific classification. Bulic & Miric (1967) reported five subspecies for the Western and Central Balkans: Citellus (—Spermophilus) citellus citellus (Linnaeus, 1766) from eastern Slavonia and Vojvodina, C. c. gradojevici Martino & Martino, 1929 from southern Macedonia, C. c. karamani Martino & Martino, 1940 from the Mt. Karadzica in
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Macedonia, C. c. Iciskarevi Martino & Martino, 1940 from Dolovo in southeastern Banat, and C. c. balcanicus Markov, 1957 from eastern Serbia. Krystufek (1990, 1996, 1998) and Krystufek & Hrabe (1996) confirmed the differences between these groups of populations on the basis of cranial measurements, variations in the pelvis (os coxae) and baculum (os penis). Only isolated mountain population (S. c. karamani) from the Mt. Jakupica and Mt. Karadžica in Macedonia is morphologically distinct in categorical criteria. The complexity of phenetic relations are shown in Fig. 18 and 19 (after Krystufek, 1996). Unfortunately, molecular analysis has not yet been performed. The research of fleas specific for S. citellus shows some similarities in geographic distribution between hosts and parasites.
Three subspecies of N. setosa are described. The range of N. s. setosa (Wagner, 1898) extends from Tien Shan in southwestern China across Kazakhstan to southwestern Russia and the Caucasus. Their major host is S. pygmaeus. Neopsylla s. moravica Rosicky, 1965 occurs in Moravia and Slovakia; its major host is S. citellus. Neopsylla s. spinea Rothschild, 1915 occupies Armenia, Romania, Bulgaria, and Yugoslavia. A major host is also S. citellus. Wagner (1939) reported this species from southern Serbia but without the exact locality, making it impossible to use these data. We provide four new localities from Yugoslavia, all located in Vojvodina: Jazak and Cortanovci from Fruška Gora in Srem, and Deliblato and Čoka from Deliblatska Peščara. There are no data for Serbia and Macedonia but they are probably distributed there because they are present in Bulgaria and Greece. The species is only rarely found on fur; it mainly lives in host nests. We examined only the fur of collected sousliks and not nests, therefore our Central Balkan findings are limited. The species is probably more common than we realize and probably occupies the entire host area.
Clenophthalmus orientalis is distributed over the entire range of S. citellus and in the Ukraine to the Ckalsky steppe ( = Čkalovnow Orenburg) in southeastern European Russia, where it parasitizes other souslik species. The nominate subspecies Ct. o. orientalis lives in the entire distribution area of S. citellus. A newly described subspecies, Ct. o. jakupicae, is found only in isolated mountain population (5. c. karamani) from the Mt. Jakupica and Mt. Karadžica in Macedonia. This agrees with earlier taxonomic conclusions (Krystufek, 1990, 1996, 1998; Krystufek & Hrabe, 1996) who considered these this population of S. citellus as morphologically most distinct from other populations.
The geographic distribution of both species from the genus Citellophilus is more complex than for N. selosa or Ct. orientalis. Both species are allopatric. Cilellophilus simplex is distributed in the eastern part of the range of S. citellus while C. martinoi mainly occupies the western part. In the central region is a mosaic of small and large populations, but the two are never simpatric. This distribution is shown on Figs. 16 and 20.
Both, C. simplex and C. martinoi, are highly variable species. Intrapopulation variation was as great as that between different populations. Variations pertain to the shape of the fixed and movable process of the clasper. Cyprich (1989) found bigger similarities in geographically neighboring than in remote populations, but they were not on the subspecific level. From both species, C. simplex and C. martinoi, is described one subspecies, which is not sharply separated from the closest populations of nominate
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Fig. 18: Geografical variations in Morphological Principal Component 1 of 16 Spennophilus citellus localities. The values are proportional to the black area in the pie diagrams, the higher the score, the darker the circle. Open circles represent the smallest sousliks, filled circles represent the largest. Male scores are on the right and female scores are on the left side of individual circles (according to Krystufek, 1966). The approximate range of the species (shaded) follows Ruzic (1978). The geographic samples are listed in Krystufek (1966).
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1.0 I_
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Fig. 19: Consensus tree summarizing ATD matrices for 16 samples of male and female Spevmophilus cite Ulis (according to Krystufek, 1966). For sample designation see Fig. 18. ATD - Average taxonomic distance, based on 13 size-adjusted cranial variables.
subspecies. Citellophilus m. rotunclus Rosicky, 1956 is restricted to the Czech Republic, Moravia, eastern Austria, western Hungary, and southern Slovakia, while C. s. rosidcy Cyprich, 1989 is found in eastern Slovakia (Fig. 20). According to Cyprich (1989) all other populations of both species from the entire range are the nominate subspecies. Cyprich was not familiar with the material from the Mt. Jakupica, where the population is morphologically different from the nominate subspecies. On the basis of 6 males, with narrower and longer fixed and narrower movable processes of the clasper (Fig. 12b), it is not possible to determine whether this material should be recognized as an independent subspecies.
Based on the distribution of C. simplex and C. martinoi and findings on the variability of citellus (Krystufek, 1990, 1996, 1998; Krystufek & Hrabe, 1996) (Fig. 18 and 19), it may be possible to determine postglacial migrations of S. citellus in Central and Eastern Europe. Further conclusions will be possible when molecular analysis of S. citellus is performed. They most probably spread in two directions: from the south towards the north and northwest and from the east towards the west. The first group, infested with C. martinoi, from central Bulgaria and western Serbia dispersed north to Romania and northwest to Vojvodina, Hungary, Austria, the Czech Republic, Moravia, and Slovakia. The second group is infested with C. simplex and spread from southern Ukraine to eastern Slovakia, eastern parts of the Pannonian lowland and to northern, eastern, and southern Bulgaria.
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Fig. 20: Southeastern Europe with survey sites of Citellophilus martinoi (Wagner & Ioff) and Citellophilus simplex (Wagner) (adapted from Cyprich, 1989 and supplemented with new findings of Brelih and Trilar).
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Acknowledgements
We would like to thank to academician Professor Bohumir Rosicky, Praha, Czech Republic; for donated material, advises and comments. Dr. Georg Dzukic, Belgrade, Yugoslavia; Dr. Boris Krystufek, Ljubljana, Slovenia; Dr. Miroljub Milenkovic, Belgrade, Yugoslavia; Dr. Boris Petrov, Belgrade, Yugoslavia; Dr. Anka Ruzic, Belgrade, Yugoslavia; and Dr. Nikola Tvrtkovic, Zagreb, Croatia, are thanked for provided material and useful comments. We are very grateful for improvements of our English text and other suggestions by Professor Jim E. Platz (Omaha, Nebraska). The research in Slovenia was partly supported by the Ministry of Science and Technology of the Republic of Slovenia (project J1-7409).
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Received I Prejeto: 12. 7. 2000
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