[Monographiae Biologicae] Biogeography and Ecology of Bulgaria Volume 82 || Fauna and Zoogeography...
Transcript of [Monographiae Biologicae] Biogeography and Ecology of Bulgaria Volume 82 || Fauna and Zoogeography...
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8 Fauna and Zoogeography of the OrthopteridInsects (Embioptera, Dermaptera,Mantodea, Blattodea, Isoptera,and Orthoptera) in Bulgaria
Alexi PopovNational Museum of Natural History, Bulgarian Academy of Sciences, Tsar Osvoboditel Blvd. 1,1000 Sofia, Bulgaria, e-mail: [email protected]
Abstract: Species diversity of orthopterid orders in Bulgaria is analyzed and revised. Four speciesand one subspecies are deleted from the list of Bulgarian fauna. Platycleis medvedeviand Eupholidoptera chabrieri are added. The number of orthopterid taxa established withcertainty in Bulgaria is 251 (238 species and 13 additional subspecies). Presence of other18 taxa (13 species and five additional subspecies) is not yet confirmed. Horizontal andvertical distribution of all species is analyzed for the first time. The richest in speciesare the Western Rhodopes (50% of the Bulgarian taxa). Characteristic species of nineregions are listed, and the fauna of these regions is compared. Of the six altitudinalbelts, the richest is the fauna of the xerothermic oak forests with 202 taxa (81 taxain this belt only). The boundaries of significant importance are the timberline and theboundary between the mesophilous forest and the beech forest. An original scheme ofzoogeographical placement of all taxa by chorotypes and by the origin is proposed. Thereare 49 types of chorotypes (geographical ranges). Most numerous are the Eurosiberianspecies (33 species, or 12.2%) but the thermophilous species (several categories) prevailin the Bulgarian fauna. Except for two oreotundral species with Arctoalpine distribution,all other taxa belong to the Arboreal type, which is divided into 23 categories according tothe origin. The Holomediterranean elements are the most numerous (37 species, or 14%).There are twice as many species with the southern (Mediterranean or Balkan) origin asthose with northern origin. The origin of southern species is analyzed for six levels ofspeciation. The most interesting zoogeographically 11 genera are analyzed in detail.
1 Introduction
The species of the order Orthoptera found in Bulgaria are well-suited for zoogeo-graphical conclusions due to several reasons. Many taxa from the superfamilyTettigonioidea and from the family Pamphagidae have restricted ranges within theBalkan Peninsula. The degree of endemism in Orthoptera is very high compared toother insect orders. Species with the southern origin prevail in the fauna of SouthEurope and the Mediterranean. These areas (including Bulgaria) are characterizedby the rich species diversity as a result of the intensive speciation in the BalkanPeninsula and Anatolia.
The position of Bulgaria at the border between the Eurosiberian and the Mediter-ranean zoogeographical subregions is the reason for its great species richness; in the
233
V. Fet and A. Popov (Eds.), Biogeography and Ecology of Bulgaria, 233–295.© 2007 Springer.
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234 A. POPOV
past, the species of both northern and southern origin dispersed into this territoryand mixed with the autochthonous fauna. The varied relief of Bulgaria with altitudesfrom the sea level up to 2925 m (Musala Peak in the Rila Mts.) is also a prerequisitefor the presence of a rich faunistic diversity of such groups as Orthoptera, whichinhabit all altitudinal vegetation belts from the xerothermic oak belt to the alpinebelt. The orography of Bulgaria and the adjacent areas has favored isolation ofcertain populations and their differentiation into distinct taxa.
The faunistic knowledge on Orthoptera and related orders in Bulgaria wasregarded as very good 15 years ago, both with respect to the species recorded inthis country as a whole and to their distribution in separate regions, as the resultof investigations by Georgi Peshev. The advances in the taxonomy in the lastperiod, however, necessitate revision of some taxa. Regarding the zoogeographicalcategorization, such categories of origin as Paleoaegeidean, Paleomediterranean,Paleoeuropean, Angaridean have been used previously (Peshev and Djingova,1974; Pešev, 1974; Pechev, 1975; Peshev and Andreeva, 1986). In general, thesecategories are quite clear but the results cannot be always compared in theoreticalreviews with the data for other insect groups. The better knowledge of the species’ranges also leads to the revision of their zoogeographical identity. It is thereforeurgent to update the faunistic data and to conduct a new zoogeographical analysisof Orthoptera, as well as related orders with lower species richness. The existingknowledge of Bulgarian fauna gives an opportunity for zoogeographical conclusionsconcerning this group of insects.
2 Faunistic Diversity
2·1 Species diversity
A list of all species recorded from Bulgaria, which have not been deleted in furtherpublications, has been compiled (Table 1). Two species newly established for thiscountry, specimens of which were found in Bulgarian collections, are also includedin this list. In total, 274 taxa (255 species and 19 additional subspecies) are listedin Table 1.
We analyzed the state of taxonomic knowledge with respect to subspecificdivision considering the possibility that some subspecies can be in fact onlyindividual aberrations or ecological forms. Subspecies of widespread specieswere not listed because of vagueness on whether they represent distinct taxa,because of the necessity of revision of subspecies, species, or groups of species(e.g. biguttulus-group of the genus Chorthippus), and because of preferred zoogeo-graphical comparison of species instead of subspecies with insufficiently knownranges. An exception is made for some verified subspecies, e.g. Platycleis albop-unctata grisea, Eupholidoptera chabrieri schmidti, Stenonemobius bicolor ponticus,Stenobothrus stigmaticus faberi as well as the endemic and subendemic subspecies,which are important for zoogeographical deductions.
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ORTHOPTERIDS OF BULGARIA 235
Tab
le1
Spec
ies
dive
rsity
,dis
trib
utio
nin
sele
cted
regi
ons,
vert
ical
dist
ribu
tion
and
zoog
eogr
aphy
ofO
rtho
pter
ida
inB
ulga
ria
Tax
aD
istr
ibut
ion
inB
ulga
ria
Veg
etat
iona
lbe
ltsin
Bul
gari
aZ
ooge
ogra
phy
W-S
TA
C-S
TA
RIL
AW
-RH
OE
-RH
OB
E,S
LB
SEA
STR
UX
ER
OA
KB
EE
CO
NSU
BA
LP
Cho
roty
peO
rigi
n
EM
BIO
PT
ER
AO
LIG
OT
OM
IDA
E
Hap
loem
bia
soli
eri
(Ram
bur,
1842
)+
++
HoM
eH
oMe/
s
DE
RM
AP
TE
RA
AN
ISO
LA
BID
IDA
E
Car
cino
phor
inae
Ani
sola
bis
mar
itim
a(G
éné,
1832
)+
+C
osH
oMe/
eS P
ON
GIP
HO
RID
AE
Lab
iinae
Lab
iam
inor
(Lin
naeu
s,17
58)
++
++
Cos
?L
AB
IDU
RID
AE
Lab
idur
inae
Lab
idur
ari
pari
a(P
alla
s,17
73)
++
++
++
+C
os?
FO
RF
ICU
LID
AE
Ane
chur
inae
Ane
chur
abi
punc
tata
(Fab
rici
us,1
781)
++
++
EuS
iSi
bF
orfi
culin
aeA
pter
ygid
am
edia
(Hag
enba
ch,1
822)
++
Eu
CE
–Me
For
ficu
laau
ricu
lari
aL
inna
eus,
1758
++
++
SL+
++
++
+C
osC
Eu
For
ficu
lasm
yrne
nsis
Serv
ille,
1839
++
+E
Me
PoM
e/s
MA
NT
OD
EA
MA
NT
IDA
E
Am
elin
aeA
mel
eshe
ldre
ichi
Bru
nner
von
Wat
tenw
yl,1
882
++
BE
++
++
EM
ePo
Me/
s
cont
inue
d
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236 A. POPOV
Tab
le1
cont
inue
d
Tax
aD
istr
ibut
ion
inB
ulga
ria
Veg
etat
iona
lbe
ltsin
Bul
gari
aZ
ooge
ogra
phy
W-S
TA
C-S
TA
RIL
AW
-RH
OE
-RH
OB
E,S
LB
SEA
STR
UX
ER
OA
KB
EE
CO
NSU
BA
LP
Cho
roty
peO
rigi
n
Man
tina
eM
anti
sre
ligi
osa
(Lin
naeu
s,17
58)
++
++
++
++
++
PT–P
aPa
Tr
Iris
orat
oria
(Lin
naeu
s,17
58)
++
+H
oMe
HoM
e/s
EM
PU
SID
AE
Em
pusi
nae
Em
pusa
fasc
iata
Bru
llé,1
836
++
++
++
+E
Me
PoM
e/s
BL
AT
TO
DE
AP
OL
YP
HA
GID
AE
Pol
ypha
gina
eP
olyp
haga
aegy
ptia
ca(L
inna
eus,
1758
)+
+T
u–M
ePo
Me/
eB
LA
TT
IDA
E
Bla
ttin
aeB
latt
aor
ient
alis
Lin
naeu
s,17
58sy
nant
hrop
icin
tow
nsan
dvi
llage
sin
the
entir
eB
ulga
ria
++
Cos
PaT
rP
erip
lane
taam
eric
ana
(Lin
naeu
s,17
58)
syna
nthr
opic
inso
me
tow
nsin
Bul
gari
a+
++
Cos
PaT
rB
LA
TT
EL
LID
AE
Bla
ttel
linae
Bla
ttel
lage
rman
ica
(Lin
naeu
s,17
67)
syna
nthr
opic
into
wns
and
villa
ges
inth
een
tire
Bul
gari
a+
+C
osPa
Tr
Lob
opte
rade
cipi
ens
(Ger
mar
,181
7)+
+B
E+
++
HoM
eH
oMe/
sE
CT
OB
IID
AE
Ect
obiin
aeE
ctob
ius
(Ect
obiu
s)sy
lves
tris
(Pod
a,17
61)
++
++
++
+E
uC
Eu
Ect
obiu
s(E
ctob
ius)
vitt
iven
tris
(Cos
ta,1
847)
+?
++
++
CSE
uPo
Me/
eE
ctob
ius
(Ect
obiu
s)la
ppon
icus
(Lin
naeu
s,17
58)
++
+SL
?+
++
++
EuS
iSi
bE
ctob
ius
(Ect
obiu
s)ba
lcan
iR
amm
e,19
23+
++
++
++
++
++
CSE
uC
Eu
Ect
obiu
s(E
ctob
ius)
eryt
hron
otus
(Bur
r,18
98)
++
++
++
+C
SEu
CE
uE
ctob
ius
(Ect
obiu
s)pu
ncta
tiss
imus
Ram
me,
1923
+T
rAd
PoM
e/e
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ORTHOPTERIDS OF BULGARIA 237
Phy
llod
rom
ica
brev
ipen
nis
(Fis
cher
,185
3)+
SL+
+E
Me
PoM
e/e
Phy
llod
rom
ica
mar
gina
ta(S
chre
ber,
1781
)+
SL+
+E
Me
PoM
e/e
Phy
llod
rom
ica
carn
ioli
ca(R
amm
e,19
13)
++
+N
WB
aD
inP
hyll
odro
mic
apa
llid
a(B
runn
ervo
nW
atte
nwyl
,188
2)+
+B
a–A
n?
Ana
Phy
llod
rom
ica
suba
pter
a(R
ambu
r,18
38)
need
sco
nfir
mat
ion
??
??
SEu
HoM
e/s
ISO
PT
ER
AK
AL
OT
ER
MIT
IDA
E
Kal
oter
mes
flav
icol
lis
(Fab
rici
us,1
793)
++
HoM
eH
oMe/
sR
HIN
OT
ER
MIT
IDA
E
Het
erot
erm
itin
aeR
etic
ulit
erm
eslu
cifu
gus
(Ros
si,1
792)
++
++
+H
oMe
HoM
e/e
OR
TH
OP
TE
RA
Ens
ifer
aT
ET
TIG
ON
IOID
EA
PH
AN
ER
OP
TE
RID
AE
Pha
nero
pter
afa
lcat
a(P
oda,
1761
)+
++
?+
++
EuS
iSi
bP
hane
ropt
era
nana
Fieb
er,1
853
++
++
++
++
HoM
eH
oMe/
eT
ylop
sis
lili
foli
a(F
abri
cius
,179
3)+
++
++
++
++
HoM
eH
oMe/
sIs
ophy
abr
evip
enni
sB
runn
ervo
nW
atte
nwyl
,187
8ne
eds
conf
irm
atio
n?
CE
uC
Eu
Isop
hya
spec
iosa
(Fri
vald
szky
,186
7)+
++
++
++
++
++
EM
ePo
Me/
eIs
ophy
are
ctip
enni
sB
runn
ervo
nW
atte
nwyl
,187
8+
++
++
Ba–
An
PoM
e/s
Isop
hya
bure
schi
Pesh
ev,1
959
++
+R
i–P–
SR
i–R
hIs
ophy
agu
lae
Pesh
ev,1
981
+T
unT
hrIs
ophy
am
odes
tior
Bru
nner
von
Wat
tenw
yl,1
882
++
+C
SEu
?M
oBa
Isop
hya
obtu
saB
runn
ervo
nW
atte
nwyl
,188
2+
++
CB
aM
oBa
Isop
hya
petk
ovi
Pesh
ev,1
959
++
++
Th–
Bl
Thr
Isop
hya
mod
esta
mod
esta
(Fri
vald
szky
,186
7)do
esno
toc
cur
inB
ulga
ria
Isop
hya
mod
esta
long
icau
data
Ram
me,
1951
++
EB
aM
oeIs
ophy
apl
evne
nsis
Pesh
ev,1
985
+C
D–C
PM
oe
cont
inue
d
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238 A. POPOV
Tab
le1
cont
inue
d
Tax
aD
istr
ibut
ion
inB
ulga
ria
Veg
etat
iona
lbe
ltsin
Bul
gari
aZ
ooge
ogra
phy
W-S
TA
C-S
TA
RIL
AW
-RH
OE
-RH
OB
E,S
LB
SEA
STR
UX
ER
OA
KB
EE
CO
NSU
BA
LP
Cho
roty
peO
rigi
n
Isop
hya
kisi
Pesh
ev,1
981
SL+
+R
i–P–
SR
i–R
hIs
ophy
aan
dree
vae
Pesh
ev,1
981
++
Stru
Mac
Isop
hya
prav
dini
prav
dini
Pesh
ev,1
985
++
CSt
StPl
Isop
hya
prav
dini
bazy
luki
Pesh
ev,1
985
++
CSt
StPl
Isop
hya
prav
dini
adam
ovic
iPe
shev
,198
5+
ESt
StPl
Isop
hya
mik
sici
Pesh
ev,1
985
++
+W
StSt
PlIs
ophy
arh
odop
ensi
sR
amm
e,19
51+
++
++
WR
hR
i–R
hIs
ophy
aho
spod
arho
spod
ar(S
auss
ure,
1898
)+
+T
h–B
lT
hrIs
ophy
aho
spod
arm
edim
onta
naN
edel
kov,
1908
+So
Mo
StPl
Isop
hya
ram
mei
Pesh
ev,1
981
+St
raT
hrB
arbi
tist
esse
rric
auda
(Fab
rici
us,1
798)
++
++
+C
SEu
CE
uB
arbi
tist
esco
nstr
ictu
sB
runn
ervo
nW
atte
nwyl
,18
78+
+C
Eu
CE
u
Anc
istr
ura
nigr
ovit
tata
(Bru
nner
von
Wat
tenw
yl,
1878
)+
++
++
++
EB
a?
Thr
Met
apla
stes
pulc
hrip
enni
s(C
osta
,186
3)do
esno
toc
cur
inB
ulga
ria
And
rein
iim
onnu
ptia
lis
(Kar
ny,1
918)
++
TrA
dM
acL
epto
phye
spu
ncta
tiss
ima
(Bos
c,17
92)
++
++
++
Eu
CE
uL
epto
phye
sal
bovi
ttat
a(K
olla
r,18
33)
++
++
BE
++
++
Eu
PoM
e/e
Lep
toph
yes
disc
oida
lis
(Fri
vald
szky
,186
7)+
++
CSE
uC
Eu
Lep
toph
yes
lati
caud
a(F
riva
ldsz
ky,1
867)
+T
rAd
Din
Pol
ysar
cus
dent
icau
da(C
harp
entie
r,18
25)
++
++
++
++
++
CSE
uC
Eu
Poe
cili
mon
orna
tus
(Sch
mid
t,18
49)
+N
WB
aD
inP
oeci
lim
onaf
fini
saf
fini
s(F
riva
ldsz
ky,1
867)
+?
+C
SEu
MoB
aP
oeci
lim
onaf
fini
sru
enen
sis
Pesh
ev,1
980
++
Oso
Ri–
Rh
Poe
cili
mon
affi
nis
rile
nsis
Pesh
ev,1
980
++
+R
i–P–
SR
i–R
hP
oeci
lim
onaf
fini
sm
edim
onta
nus
Pesh
ev,1
980
+C
SrSt
PlP
oeci
lim
onha
rzi
Pesh
ev,1
980
++
Ri–
P–S
Ri–
Rh
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ORTHOPTERIDS OF BULGARIA 239
Poe
cili
mon
mis
tshe
nkoi
mis
tshe
nkoi
Pesh
ev,1
980
+R
i–P–
SR
i–R
hP
oeci
lim
onm
ists
henk
oim
arza
niPe
shev
,198
0SL
+R
i–P–
SR
i–R
hP
oeci
lim
onm
ists
henk
oiti
nkae
Pesh
ev,1
980
BE
+B
elR
i–R
hP
oeci
lim
onm
ists
henk
oivl
ahin
ensi
sPe
shev
,198
0+
Vl–
M–O
Ri–
Rh
Poe
cili
mon
kisi
Pesh
ev,1
980
BE
+B
elR
i–R
hP
oeci
lim
onsc
hmid
ti(F
iebe
r,18
53)
++
++
++
+C
SEu
PoM
e/e
Poe
cili
mon
thor
acic
us(F
iebe
r,18
53)
++
++
++
++
++
++
+?
NC
Ba
?M
oBa
Poe
cili
mon
eleg
ans
Bru
nner
von
Wat
tenw
yl,1
878
++
++
++
+N
WB
aD
inP
oeci
lim
onfu
ssi
Bru
nner
von
Wat
tenw
yl,1
878
++
++
++
++
+C
SEu
CE
uP
oeci
lim
onzw
icki
Ram
me,
1939
++
++
++
+M
–Ri–
Rh
Mac
Poe
cili
mon
pech
evi
And
reev
a,19
78+
Vl–
M–O
Ri–
Rh
Poe
cili
mon
mac
edon
icus
Ram
me,
1926
doub
tful
info
rmat
ion
for
Bul
gari
a
Mac
Mac
Poe
cili
mon
brun
neri
(Fri
vald
szky
,186
7)+
++
++
++
++
++
+?
NC
Ba
?M
oBa
Poe
cili
mon
orbe
licu
sPa
ncic
,188
3?
++
++
++
M–R
i–R
hR
i–R
hP
oeci
lim
onbe
lasi
cens
isPo
pov,
1997
taxo
nom
icst
atus
uncl
ear
BE
+B
elR
i–R
h
Poe
cili
mon
mar
mar
aens
isN
askr
ecki
,199
1+
EB
aT
hrP
oeci
lim
onhe
inri
chi
(Ram
me,
1951
)+
Stra
Thr
Poe
cili
mon
mir
amae
Ram
me,
1933
+B
a–A
n?
Ana
ME
CO
NE
MA
TID
AE
Mec
onem
ath
alas
sinu
m(D
eG
eer,
1773
)+
++
++
++
+E
uC
Eu
Mec
onem
am
erid
iona
leC
osta
,186
0pr
obab
lyin
trod
uced
++
CSE
uA
dMe
CO
NO
CE
PH
AL
IDA
E
Con
ocep
halin
aeC
onoc
epha
lus
hast
atus
(Cha
rpen
tier,
1825
)+
++
++
++
EM
ePo
Me/
sC
onoc
epha
lus
fusc
us(F
abri
cius
,179
3)+
++
++
Pal
Si–M
eC
onoc
epha
lus
dors
alis
(Lat
reill
e,18
04)
++
++
E–W
As
Sib
Cop
ipho
rina
eR
uspo
lia
niti
dula
(Sco
poli,
1786
)+
++
++
+A
T–P
aA
fTr
cont
inue
d
![Page 8: [Monographiae Biologicae] Biogeography and Ecology of Bulgaria Volume 82 || Fauna and Zoogeography of the Orthopterid Insects (Embioptera, Dermaptera, Mantodea, Blattodea, Isoptera,](https://reader030.fdocuments.us/reader030/viewer/2022020202/575096e61a28abbf6bceacb2/html5/thumbnails/8.jpg)
240 A. POPOV
Tab
le1
cont
inue
d
Tax
aD
istr
ibut
ion
inB
ulga
ria
Veg
etat
iona
lbe
ltsin
Bul
gari
aZ
ooge
ogra
phy
W-S
TA
C-S
TA
RIL
AW
-RH
OE
-RH
OB
E,S
LB
SEA
STR
UX
ER
OA
KB
EE
CO
NSU
BA
LP
Cho
roty
peO
rigi
n
TE
TT
IGO
NII
DA
E
Tet
tigo
niin
aeT
etti
goni
avi
ridi
ssim
a(L
inna
eus,
1758
)+
++
++
++
++
++
+Pa
lSi
–Me
Tet
tigo
nia
caud
ata
(Cha
rpen
tier,
1845
)+
++
++
+E
–WA
sPo
Me/
eT
etti
goni
aca
ntan
s(F
üssl
i,17
75)
++
++
+E
uSi
Sib
Dec
ticu
sve
rruc
ivor
us(L
inna
eus,
1758
)+
++
++
++
++
++
++
Pal
Si–M
eD
ecti
cus
albi
fron
s(F
abri
cius
,177
5)+
++
++
+H
oMe
HoM
e/s
Pla
tycl
eis
(Pla
tycl
eis)
albo
punc
tata
gris
ea(F
abri
cius
,17
81)
++
++
++
++
++
+C
SEu
CE
–Me
Pla
tycl
eis
(Pla
tycl
eis)
inte
rmed
ia(S
ervi
lle,1
839)
++
++
++
++
+H
oMe
HoM
e/s
Pla
tycl
eis
(Pla
tycl
eis)
falx
(Fab
rici
us,1
775)
does
not
occu
rin
Bul
gari
aP
laty
clei
s(P
laty
clei
s)af
fini
sFi
eber
,185
3+
++
++
++
++
Tu–
Me
HoM
e/e
Pla
tycl
eis
(Pla
tycl
eis)
esca
lera
iB
oliv
ar,1
899
?+
++
EM
ePo
Me/
sP
laty
clei
s(M
onta
na)
stri
cta
(Zel
ler,
1849
)ne
eds
conf
irm
atio
n?
??
CSE
uA
dMe
Pla
tycl
eis
(Mon
tana
)m
aced
onic
a(B
erla
ndet
Cho
pard
,19
22)
++
Mac
Mac
Pla
tycl
eis
(Mon
tana
)m
edve
devi
(Mir
am,1
927)
+SE
E–W
AC
asP
laty
clei
s(M
odes
tana
)eb
neri
(Ram
me,
1926
)B
E+
NW
Ba
Din
Pla
tycl
eis
(Tes
sell
ana)
tess
ella
ta(C
harp
entie
r,18
25)
?ne
eds
conf
irm
atio
n?
Tu–
Me
HoM
e/e
Pla
tycl
eis
(Tes
sell
ana)
veys
eli
Koç
ak,1
984
++
+E
–WA
sSi
bP
laty
clei
s(T
esse
llan
a)ni
gros
igna
ta(C
osta
,186
3)+
SL+
++
EM
ePo
Me/
sP
laty
clei
s(T
esse
llan
a)or
ina
Bur
r,18
99ne
eds
conf
irm
atio
n?
SBa
BaM
e
Pla
tycl
eis
(Tes
sell
ana)
ince
rta
Bru
nner
von
Wat
tenw
yl,1
882
++
++
++
EM
ePo
Me/
s
![Page 9: [Monographiae Biologicae] Biogeography and Ecology of Bulgaria Volume 82 || Fauna and Zoogeography of the Orthopterid Insects (Embioptera, Dermaptera, Mantodea, Blattodea, Isoptera,](https://reader030.fdocuments.us/reader030/viewer/2022020202/575096e61a28abbf6bceacb2/html5/thumbnails/9.jpg)
ORTHOPTERIDS OF BULGARIA 241
Met
riop
tera
(Met
riop
tera
)bi
colo
r(P
hilip
pi,
1830
)+
++
+E
uSi
Sib
Met
riop
tera
(Met
riop
tera
)do
mog
ledi
dom
ogle
diB
runn
ervo
nW
atte
nwyl
,188
2+
++
NC
Ba
MoB
a
Met
riop
tera
(Met
riop
tera
)do
mog
ledi
arno
ldi
Ram
me,
1933
++
++
SL+
++
+C
Ba
MoB
a
Met
riop
tera
(Met
riop
tera
)ro
esel
i(H
agen
bach
,182
2)+
++
+E
uSi
Sib
Met
riop
tera
(Met
riop
tera
)fe
dtsc
henk
oiam
biti
osa
Uva
rov,
1924
taxo
nom
icst
atus
uncl
ear
++
++
SBa
BaM
e
Met
riop
tera
(Met
riop
tera
)ob
long
icol
lis
Bru
nner
von
Wat
tenw
yl,1
882
++
+SL
++
++
+SB
aB
aMe
Met
riop
tera
(Met
riop
tera
)he
ller
iSc
hmid
t,19
98+
+W
StSt
Pl
Sepi
ana
sepi
um(Y
ersi
n,18
54)
++
++
+H
oMe
HoM
e/s
Pho
lido
pter
aap
tera
apte
ra(F
abri
cius
,179
3)?
?ne
eds
conf
irm
atio
n?
CE
uC
Eu
Pho
lido
pter
aap
tera
karn
yiE
bner
,190
8+
++
++
++
+N
CB
aD
inP
holi
dopt
era
apte
rabu
lgar
ica
Mar
an,1
953
?+
++
M–R
–BM
Mac
Pho
lido
pter
abr
evip
esR
amm
e,19
39+
Ba–
An
Thr
Pho
lido
pter
ali
ttor
alis
(Fie
ber,
1853
)+
?+
+SE
uC
Eu
Pho
lido
pter
am
aced
onic
aR
amm
e,19
28+
++
+M
acM
acP
holi
dopt
era
bure
siM
aran
,195
7ta
xono
mic
stat
usun
clea
r+
Ri–
P–S
Ri–
Rh
Pho
lido
pter
aho
berl
andt
iM
aran
,195
7+
taxo
nom
icst
atus
uncl
ear
+R
i–P–
SR
i–R
h
Pho
lido
pter
arh
odop
ensi
sM
aran
,195
3+
taxo
nom
icst
atus
uncl
ear
+R
i–P–
SR
i–R
h
Pho
lido
pter
afr
ival
dsky
i(H
erm
an,1
871)
++
++
++
++
SEE
uC
Eu
Pho
lido
pter
afa
llax
(Fis
cher
,185
3)+
++
++
++
+C
SEu
PoM
e/e
Pho
lido
pter
agr
iseo
apte
ra(D
eG
eer,
1773
)+
++
+E
uC
Eu
Pho
lido
pter
aga
nevi
Har
z,19
86+
+W
Rh
Ri–
Rh
Eup
holi
dopt
era
chab
rier
isc
hmid
ti(F
iebe
r,18
61)
+T
rAd
PoM
e/s
Eup
holi
dopt
era
mar
ani
Pesh
ev,1
960
++
Stru
Mac
cont
inue
d
![Page 10: [Monographiae Biologicae] Biogeography and Ecology of Bulgaria Volume 82 || Fauna and Zoogeography of the Orthopterid Insects (Embioptera, Dermaptera, Mantodea, Blattodea, Isoptera,](https://reader030.fdocuments.us/reader030/viewer/2022020202/575096e61a28abbf6bceacb2/html5/thumbnails/10.jpg)
242 A. POPOV
Tab
le1
cont
inue
d
Tax
aD
istr
ibut
ion
inB
ulga
ria
Veg
etat
iona
lbe
ltsin
Bul
gari
aZ
ooge
ogra
phy
W-S
TA
C-S
TA
RIL
AW
-RH
OE
-RH
OB
E,S
LB
SEA
STR
UX
ER
OA
KB
EE
CO
NSU
BA
LP
Cho
roty
peO
rigi
n
Eup
holi
dopt
era
beyb
ienk
oiPe
shev
,196
2+
++
CSt
StPl
Eup
holi
dopt
era
smyr
nens
is(B
runn
ervo
nW
atte
nwyl
,188
2)+
++
Ba–
An
Ana
Par
apho
lido
pter
aca
stan
eovi
ridi
s(B
runn
ervo
nW
atte
nwyl
,188
2)+
++
+B
a–A
nA
na
Buc
epha
lopt
era
buce
phal
a(B
runn
ervo
nW
atte
nwyl
,188
2)+
++
BE
++
++
Ba–
An
Ana
Pso
rodo
notu
sfi
eber
i(F
iebe
r,18
53)
++
++
BE
++
++
NC
Ba
MoB
aP
achy
trac
his
grac
ilis
(Bru
nner
von
Wat
tenw
yl,1
861)
++
++
++
++
++
CSE
uPo
Me/
e
Pac
hytr
achi
sfr
ater
(Bru
nner
von
Wat
tenw
yl,
1882
)ne
eds
conf
irm
atio
n?
??
CSE
uD
in
Ant
eras
tes
serb
icus
Bru
nner
von
Wat
tenw
yl,
1882
++
++
++
++
+B
a–A
nA
na
Pte
role
pis
germ
anic
a(H
erri
ch-S
chäf
fer,
1840
)+
++
++
++
++
+C
SEu
PoM
e/e
Gam
psoc
leis
glab
ra(H
erbs
t,17
86)
?+
E–W
As
Cas
Gam
psoc
leis
abbr
evia
taH
erm
an,1
874
SL+
SBa
BaM
eO
ncon
otin
aeO
ncon
otus
serv
ille
iFi
sche
rde
Wal
dhei
m,
1846
+SE
E–W
AC
as
Sagi
nae
Saga
nato
liae
Serv
ille,
1839
++
+B
E+
++
++
EM
ePo
Me/
sSa
gacf
.hel
leni
caK
alte
nbac
h,19
67+
SBa
BaM
eSa
gara
mm
eiK
alte
nbac
h,19
65+
+M
a–T
hM
acSa
gaca
mpb
elli
cam
pbel
liU
varo
v,19
21+
++
++
Mac
Mac
Saga
cam
pbel
ligr
acil
isK
is,1
962
++
+E
Ba
Thr
Saga
pedo
(Pal
las,
1771
)+
+E
–WA
sC
as
![Page 11: [Monographiae Biologicae] Biogeography and Ecology of Bulgaria Volume 82 || Fauna and Zoogeography of the Orthopterid Insects (Embioptera, Dermaptera, Mantodea, Blattodea, Isoptera,](https://reader030.fdocuments.us/reader030/viewer/2022020202/575096e61a28abbf6bceacb2/html5/thumbnails/11.jpg)
ORTHOPTERIDS OF BULGARIA 243
BR
AD
YP
OR
IDA
E
Eph
ippi
geri
nae
Eph
ippi
ger
ephi
ppig
erep
hipp
iger
(Fie
big,
1784
)+
++
++
++
++
+C
SEu
CE
u
Eph
ippi
ger
ephi
ppig
erba
lkan
icus
And
reev
a,19
85+
+ta
xono
mic
stat
usun
clea
r+
++
+N
Bu
StPl
Eph
ippi
ger
ephi
ppig
erva
rnen
sis
And
reev
a,19
85ta
xono
mic
stat
usun
clea
r+
+N
Bl
Moe
Bra
dypo
rina
eB
rady
poru
sda
sypu
s(I
llige
r,18
00)
++
++
++
+E
Ba
?M
oeC
alli
men
usm
acro
gast
er(L
efeb
vre,
1831
)+
++
+B
a–A
n?
Moe
RH
APH
IDO
PHO
RO
IDE
AR
HA
PH
IDO
PH
OR
IDA
E
Tro
glop
hilin
aeT
rogl
ophi
lus
negl
ectu
svl
asin
ensi
sM
aran
,19
58+
++
++
subs
peci
esst
atus
uncl
ear
CB
aSt
Pl
Rha
phid
opho
rina
eT
achy
cine
sas
ynam
orus
Ade
lung
,190
2ac
cide
ntal
lyin
trod
uced
inth
epa
st;
not
rece
ntly
dist
ribu
ted
Cos
SiT
i
GR
YL
LO
IDE
AG
RY
LL
IDA
E
Gry
llina
eG
ryll
usca
mpe
stri
sL
inna
eus,
1758
++
++
++
++
++
+W
PaC
E–M
eG
ryll
usbi
mac
ulat
usD
eG
eer,
1773
doub
tful
info
rmat
ion
for
Bul
gari
aA
T–P
aA
fTr
Ach
eta
dom
esti
cus
(Lin
naeu
s,17
58)
syna
nthr
opic
invi
llage
s+
++
Cos
HoM
e/e
Mel
anog
ryll
usde
sert
us(P
alla
s,17
71)
++
++
+W
PaH
oMe/
eT
arta
rogr
yllu
ssa
ndan
ski
And
reev
a,19
82+
+St
ruM
acE
umod
icog
ryll
usbo
rdig
alen
sis
(Lat
reill
e,18
04)
++
Tu–
Me
HoM
e/e
Mod
icog
ryll
usfr
onta
lis
(Fie
ber,
1844
)+
++
++
E–W
As
Cas
Mod
icog
ryll
ustr
unca
tus
(Tar
bins
ky,1
940)
++
SEE
u?
Gry
llom
orph
inae
Gry
llom
orph
ada
lmat
ina
(Ocs
kay,
1832
)+
++
HoM
eH
oMe/
sG
ryll
omor
pha
cf.m
iram
aeM
edve
dev,
1933
?+
+SE
E–W
A?
PoM
e/e
Dis
copt
ila
bure
siM
aran
,195
8+
++
ER
–Bl
Thr
cont
inue
d
![Page 12: [Monographiae Biologicae] Biogeography and Ecology of Bulgaria Volume 82 || Fauna and Zoogeography of the Orthopterid Insects (Embioptera, Dermaptera, Mantodea, Blattodea, Isoptera,](https://reader030.fdocuments.us/reader030/viewer/2022020202/575096e61a28abbf6bceacb2/html5/thumbnails/12.jpg)
244 A. POPOV
Tab
le1
cont
inue
d
Tax
aD
istr
ibut
ion
inB
ulga
ria
Veg
etat
iona
lbe
ltsin
Bul
gari
aZ
ooge
ogra
phy
W-S
TA
C-S
TA
RIL
AW
-RH
OE
-RH
OB
E,S
LB
SEA
STR
UX
ER
OA
KB
EE
CO
NSU
BA
LP
Cho
roty
peO
rigi
n
Nem
obiin
aeP
tero
nem
obiu
she
yden
ihe
yden
i(F
isch
er,1
853)
++
+?
SL+
?+
+C
SEu
HoM
e/e
Pte
rone
mob
ius
heyd
eni
tart
arus
(Sau
ssur
e,18
74)
++
SEE
–WA
?C
asSt
enon
emob
ius
bico
lor
pont
icus
Gor
ocho
v,19
84+
SEE
uPo
Me/
sO
ecan
thin
aeO
ecan
thus
pell
ucen
s(S
copo
li,17
63)
++
++
BE
++
++
WPa
HoM
e/e
MO
GO
PL
IST
IDA
E
Ara
chno
ceph
alus
vest
itus
Cos
ta,1
855
++
++
SEu
HoM
e/s
MY
RM
EC
OP
HIL
IDA
E
Myr
mec
ophi
lus
acer
voru
m(P
anze
r,[1
799]
)?
?+
CSE
uC
Eu
Myr
mec
ophi
lus
myr
mec
ophi
lus
(Sav
i,18
19)1
++
SEu
HoM
e/s
GR
YL
LO
TA
LP
IDA
E
Gry
llot
alpa
gryl
lota
lpa
(Lin
naeu
s,17
58)
s.l.
++
++
++
++
+sp
ecie
sco
mpl
exW
PaC
Eu
Cae
lifer
aT
ET
RIG
OID
EA
TE
TR
IGID
AE
Tet
rix
subu
lata
(Lin
naeu
s,17
58)
++
++
++
++
+H
olSi
–Ne
Tet
rix
boli
vari
Saul
cy,1
901
++
++
++
Tu–
Me
HoM
e/e
Tet
rix
cepe
roi
(Bol
ivar
,188
7)+
++
+W
PaH
oMe/
eT
etri
xtu
erki
(Kra
uss,
1876
)+
++
++
+C
SEu
CE
uT
etri
xbi
punc
tata
(Lin
naeu
s,17
58)
++
+B
E+
++
Pal
Si–M
eT
etri
xte
nuic
orni
s(S
ahlb
erg,
1893
)+
++
++
++
++
++
Pal
Si–M
eU
varo
vite
ttix
depr
essu
s(B
riso
utde
Bar
nevi
lle,
1848
)+
++
++
++
++
++
+T
u–M
eH
oMe/
e
Par
atet
tix
mer
idio
nali
s(R
ambu
r,18
38)
++
++
++
HoM
eH
oMe/
sT
RID
AC
TY
LO
IDE
AT
RID
AC
TY
LID
AE
Xya
vari
egat
aL
atre
ille,
1809
??
+?
?+
??
PT–P
a?
Xya
pfae
ndle
ri(H
arz,
1970
)+
+PT
–Pa
?
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ORTHOPTERIDS OF BULGARIA 245
AC
RID
OID
EA
PA
MP
HA
GID
AE
Pam
phag
inae
Par
anoc
arod
esst
raub
eist
raub
ei(F
iebe
r,18
53)
++
Ba–
An
Ana
Par
anoc
arod
esch
opar
diPe
shev
,196
5+
+E
Rh
Thr
Par
anoc
arac
ris
bulg
aric
usbu
lgar
icus
(Ebn
eret
Dre
now
ski,
1930
)+
SL+
++
++
M–R
–BM
Ri–
Rh
Aki
ceri
nae
Gly
phan
usob
tusu
sFi
eber
,185
3do
esno
tocc
urin
Bul
gari
aA
siot
met
his
lim
batu
s(C
harp
entie
r,18
45)
++
SL+
++
Ba–
An
?A
naA
CR
IDID
AE
Cat
anto
pina
eP
odis
ma
pede
stri
s(L
inna
eus,
1758
)+
++
+?
?+
++
+E
uSi
Sib
Boh
eman
ella
frig
ida
(Boh
eman
,184
6)+
SL+
+H
-OT
uSi
Tu
Mir
amel
lasp
.+
+?
CB
a?
Din
Pse
udop
odis
ma
fieb
eri
(Scu
dder
,189
7)+
++
++
SEu
MoB
aO
dont
opod
ism
asc
hmid
ti(F
iebe
r,18
53)
++
++
CSE
u?
Din
Odo
ntop
odis
ma
rubr
ipes
(Ram
me,
1931
)+
++
++
+C
SEu
CE
uO
dont
opod
ism
ade
cipi
ens
Ram
me,
1951
++
++
++
++
++
SEE
uC
Eu
Pez
otet
tix
gior
nae
(Ros
si,1
794)
++
++
++
++
++
+H
oMe
HoM
e/e
Cal
lipta
min
aeC
alli
ptam
usit
alic
us(L
inna
eus,
1758
)+
++
++
++
++
++
Tu–
Me
HoM
e/e
Cal
lipt
amus
barb
arus
(Cos
ta,1
836)
++
+B
E+
++
+T
u–M
eH
oMe/
eP
arac
alop
tenu
sca
lopt
enoi
des
(Bru
nner
von
Wat
tenw
yl,1
861)
++
++
++
++
++
++
EM
ePo
Me/
e
Cyr
taca
ntha
crid
inae
Ana
crid
ium
aegy
ptiu
m(L
inna
eus,
1764
)+
BE
++
+A
T–P
aH
oMe/
eA
crid
inae
Acr
ida
unga
rica
(Her
bst,
1786
)+
++
++
++
++
HoM
eH
oMe/
eO
edip
odin
aeL
ocus
tam
igra
tori
a(L
inna
eus,
1758
)+
++
++
PT–P
aPa
Tr
Oed
aleu
sde
coru
s(G
erm
ar,1
826)
++
++
++
++
AT
–Pa
HoM
e/e
cont
inue
d
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246 A. POPOV
Tab
le1
cont
inue
d
Tax
aD
istr
ibut
ion
inB
ulga
ria
Veg
etat
iona
lbe
ltsin
Bul
gari
aZ
ooge
ogra
phy
W-S
TA
C-S
TA
RIL
AW
-RH
OE
-RH
OB
E,S
LB
SEA
STR
UX
ER
OA
KB
EE
CO
NSU
BA
LP
Cho
roty
peO
rigi
n
Pso
phus
stri
dulu
s(L
inna
eus,
1758
)+
++
++
++
++
EuS
iSi
bC
eles
vari
abil
is(P
alla
s,17
71)
++
++
E–W
As
Cas
Oed
ipod
aca
erul
esce
nsca
erul
esce
ns(L
inna
eus,
1758
)+
++
++
++
++
++
+Pa
lSi
–Me
Oed
ipod
aca
erul
esce
nsga
nevi
Har
z,19
85st
atus
ofth
esu
bspe
cies
uncl
ear
++
++
Vl–
M–O
Ri–
Rh
Oed
ipod
age
rman
ica
(Lat
reill
e,18
04)
++
++
++
++
++
+C
SEu
HoM
e/e
Oed
ipod
am
inia
ta(P
alla
s,17
71)
++
++
BE
++
++
Tu–
Me
HoM
e/e
Sphi
ngon
otus
caer
ulan
s(L
inna
eus,
1767
)+
++
++
++
++
E–W
As
CE
uA
crot
ylus
patr
ueli
s(H
erri
ch-S
chäf
fer,
1838
)+
++
BE
++
+A
T–P
aA
fTr
Acr
otyl
usin
subr
icus
(Sco
poli,
1786
)+
++
++
++
++
+A
T–P
aA
fTr
Acr
otyl
uslo
ngip
es(C
harp
entie
r,18
45)
++
++
++
AT
–Pa
AfT
rA
iolo
pus
thal
assi
nus
(Fab
rici
us,1
781)
++
++
++
+C
osPa
Tr
Aio
lopu
sst
repe
ns(L
atre
ille,
1804
)+
++
++
++
++
++
AT
–Pa
AfT
rE
pacr
omiu
ste
rges
tinu
s(C
harp
entie
r,18
25)
++
+Pa
lM
onE
pacr
omiu
sco
erul
ipes
(Iva
nov,
1887
)+
+E
uSi
Sib
Pla
typy
gius
cras
sus
(Kar
ny,1
907)
++
SEE
uC
asM
ecos
teth
uspa
rapl
euru
s(H
agen
bach
,182
2)+
++
++
EuS
iSi
bSt
etho
phym
agr
ossu
m(L
inna
eus,
1758
)+
++
++
+E
uSi
Sib
Par
acin
ema
tric
olor
(Thu
nber
g,18
15)
++
+A
T–P
aA
fTr
Dur
onie
lla
lati
corn
is(K
raus
s,19
09)
++
EM
ePo
Me/
sG
omph
ocer
inae
Arc
ypte
ra(A
rcyp
tera
)fu
sca
(Pal
las,
1773
)+
++
++
++
EuS
iSi
bA
rcyp
tera
(Par
arcy
pter
a)m
icro
pter
a(F
isch
erde
Wal
dhei
m,1
833)
++
++
++
EuS
iSi
b
Pal
lasi
ella
turc
oman
a(F
isch
erde
Wal
dhei
m,
1833
)+
+SE
E–W
A?
PoM
e/e
Chr
ysoc
hrao
ndi
spar
(Ger
mar
,[18
34])
++
++
+E
uSi
Sib
Eut
hyst
ira
brac
hypt
era
(Ocs
kay,
1826
)+
++
++
++
+E
uSi
Sib
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ORTHOPTERIDS OF BULGARIA 247
Doc
iost
auru
sm
aroc
canu
s(T
hunb
erg,
1815
)+
++
+B
E+
++
++
Tu–
Me
HoM
e/e
Doc
iost
auru
sbr
evic
olli
s(E
vers
man
n,18
48)
++
++
++
++
++
+SE
E–W
A?
Cas
Doc
iost
auru
sge
nei
(Ocs
kay,
1833
)+
+H
oMe
HoM
e/s
Doc
iost
auru
sta
rtar
usU
varo
v,19
21ne
eds
conf
irm
atio
n?
?T
u–C
aIr
a
Doc
iost
auru
skr
auss
i(I
ngen
izki
j,18
97)
++
SEE
–WA
?C
asN
otos
taur
usan
atol
icus
(Kra
uss,
1896
)+
++
+E
Me
PoM
e/e
Om
oces
tus
viri
dulu
s(L
inna
eus,
1758
)+
++
++
++
+E
uSi
Sib
Om
oces
tus
rufi
pes
(Zet
ters
tedt
,182
1)+
++
++
++
++
++
+Pa
lSi
–Me
Om
oces
tus
min
utus
(Bru
llé,1
832)
++
+B
E+
++
+SE
Eu
PoM
e/e
Om
oces
tus
petr
aeus
(Bri
sout
deB
arne
ville
,18
55)
++
++
+B
E+
++
++
EuS
iSi
–Me
Om
oces
tus
haem
orrh
oida
lis
(Cha
rpen
tier,
1825
)+
++
++
++
++
++
+E
uSi
Sib
Sten
obot
hrus
cras
sipe
s(C
harp
entie
r,18
25)
++
++
+C
SEu
CE
uSt
enob
othr
usli
neat
us(P
anze
r,17
96)
++
++
++
++
++
EuS
iSi
bSt
enob
othr
usni
grom
acul
atus
(Her
rich
-Sch
äffe
r,18
40)
++
++
BE
?+
++
++
+E
uSi
Sib
Sten
obot
hrus
fisc
heri
(Eve
rsm
ann,
1848
)+
++
++
SL+
++
++
+E
uSi
Sib
Sten
obot
hrus
bulg
aric
usR
amm
e,19
33ta
xono
mic
stat
usun
clea
r+
+W
Rh
Ri–
Rh
Sten
obot
hrus
stig
mat
icus
fabe
riH
arz,
1975
++
++
BE
++
++
+C
SEu
CE
uSt
enob
othr
usru
bicu
ndul
usK
ruse
man
etJe
ekel
,19
67+
++
+SL
++
+C
SEu
CE
u
Gom
phoc
erus
sibi
ricu
s(L
inna
eus,
1767
)+
++
++
++
++
EuS
iSi
bG
omph
ocer
ippu
sru
fus
(Lin
naeu
s,17
58)
++
++
++
++
EuS
iSi
bA
erop
edel
lus
vari
egat
us(F
isch
erde
Wal
dhei
m,
1846
)+
++
H-O
Tu
SiT
u
Myr
mel
eote
ttix
mac
ulat
us(T
hunb
erg,
1815
)+
++
++
?+
++
+E
uSi
Sib
Stau
rode
rus
scal
aris
(Fis
cher
deW
aldh
eim
,18
46)
++
++
++
++
+E
uSi
Sib
Cho
rthi
ppus
vaga
ns(E
vers
man
n,18
48)
+?
?+
CSE
uC
Eu
Cho
rthi
ppus
mol
lis
mol
lis
(Cha
rpen
tier,
1825
)+
++
++
++
++
EuS
iSi
bC
hort
hipp
usm
olli
spe
chev
iK
aram
an,1
975
stat
usof
the
subs
peci
esun
clea
r
++
NB
lM
oe
cont
inue
d
![Page 16: [Monographiae Biologicae] Biogeography and Ecology of Bulgaria Volume 82 || Fauna and Zoogeography of the Orthopterid Insects (Embioptera, Dermaptera, Mantodea, Blattodea, Isoptera,](https://reader030.fdocuments.us/reader030/viewer/2022020202/575096e61a28abbf6bceacb2/html5/thumbnails/16.jpg)
248 A. POPOV
Tab
le1
cont
inue
d
Tax
aD
istr
ibut
ion
inB
ulga
ria
Veg
etat
iona
lbe
ltsin
Bul
gari
aZ
ooge
ogra
phy
W-S
TA
C-S
TA
RIL
AW
-RH
OE
-RH
OB
E,S
LB
SEA
STR
UX
ER
OA
KB
EE
CO
NSU
BA
LP
Cho
roty
peO
rigi
n
Cho
rthi
ppus
brun
neus
(Thu
nber
g,18
15)
++
++
++
++
++
++
+E
uSi
Sib
Cho
rthi
ppus
born
halm
iH
arz,
1971
++
++
EM
ePo
Me/
sC
hort
hipp
usbi
gutt
ulus
(Lin
naeu
s,17
58)
++
++
++
++
++
++
Pal
Sib
Cho
rthi
ppus
porp
hyro
pter
useu
hedi
ckei
Hel
vers
en,1
989
++
Ba–
An
BaM
e
Cho
rthi
ppus
apri
cari
us(L
inna
eus,
1758
)+
++
++
++
+E
uSi
Sib
Cho
rthi
ppus
para
llel
uspa
rall
elus
(Zet
ters
tedt
,18
21)
++
++
+B
E+
++
++
++
EuS
iSi
b
Cho
rthi
ppus
para
llel
uste
nuis
(Bru
llé,1
832)
+B
a–A
nB
aMe
Cho
rthi
ppus
mon
tanu
s(C
harp
entie
r,18
25)
++
+E
uSi
Sib
Cho
rthi
ppus
albo
mar
gina
tus
(De
Gee
r,17
73)
++
++
++
++
++
EuS
iSi
bC
hort
hipp
usdo
rsat
us(Z
ette
rste
dt,1
821)
++
++
++
++
EuS
iSi
bC
hort
hipp
usdi
chro
us(E
vers
man
n,18
59)
++
++
++
EuS
iSi
bC
hort
hipp
uslo
ratu
s(F
isch
erde
Wal
dhei
m,
1846
)+
++
++
+E
Me
Cas
Euc
hort
hipp
uspu
lvin
atus
(Fis
cher
deW
aldh
eim
,184
6)+
++
++
++
Tu–
Me
Cas
Euc
hort
hipp
usde
cliv
us(B
riso
utde
Bar
nevi
lle,
1848
)+
++
++
++
++
++
CSE
uH
oMe/
e
1D
urin
gth
epr
intin
gof
the
pres
ent
pape
r,th
eon
lysp
ecim
enpu
blis
hed
asM
yrm
ecop
hilu
sm
yrm
ecop
hilu
sfr
omB
ulga
ria
(Pop
ovan
dC
hoba
nov,
2004
)w
asid
entif
ied
byD
r.A
.Gor
ocho
vas
Myr
mec
ophi
lus
hirt
icau
dus
(Fis
cher
deW
aldh
eim
,184
6).T
hefi
ndin
gof
mor
em
ater
ial
will
solv
eth
epr
oble
mto
whi
chsp
ecie
sbe
long
sth
issp
ecim
en.
Reg
ions
ofB
ulga
ria:
BE
,B
elas
itsa
Mts
.;B
SEA
,B
lack
Sea
coas
t;C
-ST
A,
Cen
tral
Star
aPl
anin
aN
atio
nal
Park
;E
-RH
O,
Eas
tern
Rho
dope
sM
ts.;
RIL
A,
Rila
Nat
iona
lPa
rk;
SL,S
lavy
anka
Mts
.;ST
RU
,Mid
dle
Stru
ma
Val
ley;
W-R
HO
,Wes
tern
Rho
dope
sM
ts.;
W-S
TA
,Wes
tern
Star
aPl
anin
aM
ts.
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ORTHOPTERIDS OF BULGARIA 249
Veg
etat
iona
lbe
lts:
AL
P,al
pine
;B
EE
,bee
ch;
CO
N,c
onif
erou
s;O
AK
,mes
ophi
lous
oak;
SUB
,sub
alpi
ne;
XE
R,x
erot
herm
icoa
k.C
horo
type
s:A
T–P
a,A
frot
ropi
cal–
Pale
arct
ic;B
a–A
n,B
alka
n–A
nato
lian;
Bel
,Bel
asits
a;C
Ba,
Cen
tral
Bal
kan;
CD
–CP,
Cen
tral
Dan
ubia
n–C
entr
alPr
edba
lkan
;CE
u,C
entr
alE
urop
ean;
Cos
,C
osm
opol
itan;
CSE
u,C
entr
alan
dSo
uth
Eur
opea
n;C
Sr,
Cen
tral
Sred
naG
ora;
CSt
,C
entr
alSt
ara
Plan
ina;
EB
a,E
aste
rnB
alka
n;E
Me,
Eas
tern
Med
iterr
anea
n;E
R–B
l,E
aste
rnR
hodo
pean
–Bla
ckSe
aco
ast;
ER
h,E
aste
rnR
hodo
pean
;E
St,
Eas
tern
Star
aPl
anin
a;E
u,E
urop
ean;
EuS
i,E
uros
iber
ian;
E–W
As,
Eur
opea
n–W
este
rnA
sian
;H
ol,H
olar
ctic
;H
oMe,
Hol
omed
iterr
anea
n;H
-OT
u,H
olar
ctic
Ore
otun
dral
;M
ac,M
aced
onia
n;M
a–T
h,M
aced
onia
n–T
hrac
ian;
M–R
–BM
,Mac
edon
ian–
Rho
dope
an–B
lack
orM
arm
ara
Sea
coas
t;M
–Ri–
Rh,
Mac
edon
ian–
Rila
–Rho
dope
an;N
Bl,
nort
hern
part
ofB
ulga
rian
Bla
ckSe
aco
ast;
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250 A. POPOV
The occurrence of each species in Bulgaria was also analyzed. The presence ofmaterial was checked in the collections of the National Museum of Natural Historyin Sofia (NMNHS) where the collection of G. Peshev is deposited. The correctnessof interpretation of the published species records based on material which has notbeen preserved was estimated depending on the time of identification and the useof literature relevant from the current point of view.
Deleted taxa. Of the taxa given in Table 1, three species and one subspecies,whose occurrence in Bulgaria is out of the question, as well as one species, whichhad been introduced accidentally in the past but does not occur now in this country,have to be deleted from the list of Bulgarian fauna. Metaplastes pulchripennis,Platycleis falx, and Isophya modesta modesta (only as a subspecies) have been incor-rectly identified. They are not found in the Balkan Peninsula, and the informationfor Bulgaria must be rejected for zoogeographical reasons. Glyphanus obtusus isan endemic species for the eastern part of Central Greece and the adjacent areasof Peloponnesus. The statement that the terra typica of the species is situated inBulgaria is a result of an erroneous interpretation of the locality. The differentcase is Tachycines asynamorus, which was accidentally introduced in greenhousesin Sofia where it has occurred for 15 years. During the last 60 years this specieshas not been found in Bulgaria and should be deleted from the list of Bulgarianfauna. Along with the deleted taxa, the subfamily Rhaphidophorinae and thegenera Metaplastes (for the present), Glyphanus, and Tachycines have to be deletedas well.
Added taxa. Two newly recorded species, which have not been published sofar from Bulgaria, are Platycleis (Montana) medvedevi from North Bulgaria andEupholidoptera chabrieri (with its subsp. schmidti) from Southwestern Bulgaria(Fig. 4).
Number of taxa. With the abovementioned corrections, the total number oftaxa of Orthopterida in Bulgaria is 269, including 251 species and 18 additionalsubspecies. They are distributed by orders as follows: Embioptera, one species;Dermaptera, seven species; Mantodea, four species; Blattodea, 16 species; Isoptera,two species; Orthoptera, 221 species and 18 subspecies, or 239 taxa. This numberincludes 238 species and 13 additional subspecies, or 251 taxa altogether, whichare recorded with certainty for the fauna of Bulgaria. The remaining taxa publishedfor Bulgaria cannot be confirmed or have unclear taxonomic status. Their occur-rence in Bulgaria, however, is possible; they are retained in the list until theywill be confirmed or rejected, or until their status as distinct taxa will beclarified. Such taxa, for which occurrence in Bulgaria is doubtful, belong to threecategories.
Species with doubtful locality data for Bulgaria. These are Poecilimonmacedonicus and Gryllus bimaculatus, published for Bulgaria without exact local-ities and most likely reported as a speculation and not based of a concreterecord.
Taxa needing confirmation: eight taxa (seven species and one subspecies).Blattodea and Caelifera are represented with one species each; the remaining taxa
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ORTHOPTERIDS OF BULGARIA 251
belong to the superfamily Tettigonioidea. The doubt in their correct identificationarises due to lack of confirmation of the records dated 100 years ago (Platycleisstricta) or 50 years ago (Platycleis tessellata and Pachytrachis frater), from therecord based only on a female (Isophya brevipennis), or from identification thatseems uncertain from the current point of view (Phyllodromica subaptera fromBlattodea, Platycleis orina, Dociostaurus tartarus, and Pholidoptera aptera apteraas a subspecies).
Taxa with unclear status. Possible synonyms are four species of Pholidoptera andStenobothrus as well as four subspecies of Ephippiger, Oedipoda, and Chorthippus,all described from Bulgaria. This, however, must be proved through taxonomicinvestigations.
The next two categories include species which are a part of the Bulgarian fauna,though not autochthonous.
Introduced species. The only species accidentally introduced to Bulgaria isMeconema meridionale. A small population has been found recently by Chobanov(2003) in a Botanical Garden north of Varna. At the same place and timeChobanov (2003) found also small but stable introduced population of Leptophyespunctatissima; however, this species occur autochthonously in Western Bulgaria. Inthe same paper, MSc Dragan Chobanov proposes two hypotheses for the occurrenceof Barbitistes constrictus in the Rila Mts. One of these considers this species as intro-duced accidentally in the beginning of the 20th century. I regard as more probablethe alternative hypothesis about a relict origin of the Rila population. ConcerningTachycines asynamorus, accidentally introduced 70 years ago but extinct later, seeabove under Deleted taxa.
Synanthropic species. Three species of Blattodea and one species of Grylloideabelong to this category. Bulgaria has not been a part of the original range of thesespecies. This is indisputable for the representatives of Blattodea and very likely forAcheta domesticus. It is possible that the native area of the latter is located near toBulgaria. Nevertheless, it seems that this cricket species was also spread passivelybecause (as well as three cockroache species) it does not occur in Bulgaria outsideof the houses.
2·2 Taxonomic and faunistic problems
The advances of the orthopterid taxonomy in the last 20 years led to thestatus changes for some taxa. New species and subspecies were described,new synonyms established, some species split, and the status of some taxaelevated or downgraded. Nearly all published faunistic information for Bulgariapredates this period. Also, part of it had not been interpreted according to theknowledge available at the time of publication. Because of this, the accumu-lation of taxonomic problems concerning the species diversity in Bulgaria, whichawait the solution, is not surprising. These problems can be summarized asfollows.
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252 A. POPOV
Taxonomic problems
1. Validity of taxa described from Bulgaria. Many species and subspecies fromthe genera Isophya, Poecilimon, Eupholidoptera, and Ephippiger, described byG. Peshev, E. Andreeva, and N. Nedelkov, need to be revised. Only few of themare likely to be distinct species.
2. Probable synonyms. New material of Stenobothrus bulgaricus is necessary to befound in its type locality. Poecilimon belasicensis needs to be compared withrelated species from adjacent countries.
3. Revision of species groups. The aptera-group of the genus Pholidoptera needs arevision. Three species described by Dr. Josef Maran are very likely synonymsbut due to the small number of known specimens it is not clear to which speciesthey belong. The revision will show the systematic position of the taxon oftenrecorded from South Bulgaria and reported as Pholidoptera sp. This is possiblyone of the three subspecies of Pholidoptera aptera or Ph. macedonica found inSouth Bulgaria and North Greece.
4. Verification of the taxonomic status. It has to be checked whether Metri-optera domogledi domogledi and M. d. arnoldi occur sympatrically on StaraPlanina; in this case they are distinct species. They were regarded for along time as species with distinct differences in the subgenital plate inboth sexes but have been downgraded to subspecies status because of thesimilarity in the song pattern. Another problem in the same genus concernsM. roeseli and M. fedtschenkoi ambitiosa. Recently, a doubt was expressedabout the occurrence of the latter along with the former in the BalkanPeninsula. A question arises about what are the populations published asM. f. ambitiosa.
5. Unclear status of subspecies. The distinct subspecies status of Troglophilusneglectus vlasinensis, Chorthippus parallelus tenuis, Ch. mollis pechevi, andOedipoda caerulescens ganevi has to be verified. The last two subspecies aremost likely synonyms but this has to be proved. The subspecific division ofSaga campbelli and Callimenus macrogaster also awaits revision.
6. Unknown male. For Pholidoptera ganevi, it is not possible to determine itssystematic position in the genus and the species validity until the male will bedescribed.
Faunistic problems
7. Species diversity of genera and subgenera. The genera Ectobius (Blattodea)and Pachytrachis and the subgenera Montana and Tessellana (both of the genusPlatycleis) in Bulgaria need a revision; it has to be established which, andhow many, species of each of these genus-group taxa occur in Bulgaria. Thepublished species of Ectobius have not been identified according to the current
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ORTHOPTERIDS OF BULGARIA 253
criteria (a glandular pit on the seventh tergite in males). Attention should bepaid particularly to the complex Ectobius erythronotus / E. burri.
8. Genus known for Bulgaria from females only. The genus Miramella wasreported for Bulgaria only once (Nedelkov, 1908) as Miramella alpina, the onlyspecies of the genus known at that time. Only females, which are unidentifiableunder the current criteria, have been found. The species is now split in manyspecies and the occurrence of four of them in Bulgaria is possible. The issuecannot be settled until the male is found.
9. Unknown or unclear subspecies identity. The material of Pholidopteramacedonica and Callimenus macrogaster has to be revised in order to showwhich, and how many, of the subspecies of these two species occur inBulgaria. Three subspecies of Pholidoptera aptera are reported from Bulgaria.Ph. a. karnyi is common in the mountains (Fig. 6), and Ph. a. bulgarica is verylikely also a distinct subspecies distributed at least in the lowlands. The recordof Ph. a. aptera in Bulgaria seems dubious.
10. Insufficiently known distribution in Bulgaria of closely related species. Speciesof the genera Modicogryllus and Xya have been split in two species each butreported for Bulgaria on the basis of outdated taxonomic criteria. Along withModicogryllus frontalis and Xya variegata, the closely related M. truncatusand X. pfaendleri are recorded in Bulgaria as well. The correct identificationwill eliminate the confusion and will outline the distribution of each species inBulgaria.
11. Species whose identification is not clear. The recent records of Sagacf. hellenica and Gryllomorpha cf. miramae (Chobanov, 2003) needs verifi-cation on the basis of more abundant material from Bulgaria and comparativematerial from other countries.
12. Doubtful records. Two species, Poecilimon macedonicus and Gryllus bimacu-latus, reported for Bulgaria (most likely on the basis of speculations), have tobe searched both in collections and in nature.
13. Probable incorrect identification. Three species, Phyllodromica subaptera(Blattodea), Isophya brevipennis, and Dociostaurus tartarus, have to besearched in collections or in the reported localities. The report of Ph. subapterais most likely an incorrect identification but the occurrence of this species inBulgaria is possible. The last two species are not very likely to be found inBulgaria.
14. Stability of the population of an introduced species. Future studies will showwhether the small introduced population of Meconema meridionale in theBotanical Garden near Varna (Chobanov, 2003) will survive and whether itwill expand from this locality (the only in Bulgaria).
15. Sibling species distinguished genetically. Gryllotalpa gryllotalpa is considerednow a complex of species with a different number of chromosomes. Theirtaxonomic status in most cases is not clarified. Whether one or more species,and which ones exactly, occur in Bulgaria can be decided only after geneticstudies.
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254 A. POPOV
3 Distribution of Species
3·1 Horizontal distribution
Nine regions with well, and relatively evenly, explored fauna of Orthoptera areselected to illustrate its distribution in Bulgaria. One or two reviews for each ofthese regions have been published in the last five years (Central Stara PlaninaNational Park1, Rila National Park, Kresna Gorge, Eastern Rhodopes Mts.) or 30–40years earlier (Belasitsa Mts., Slavyanka Mts., Western Stara Planina Mts., BlackSea coast, Western Rhodopes).
These regions have been selected to cover the maximal difference in theecological conditions which is the premise for variation in faunistic diversity. Thenine selected regions house 91% of the orthopterid species in Bulgaria (Table 1).The majority of the remaining 23 species occur in Pirin and Strandja; these mountainranges are not included since they are insufficiently studied. The selected regionsinclude the high mountain ranges with coldest and most humid climate (Rila,Stara Planina, Western Rhodopes) and the three regions with warmest and driestclimate in this country: the Middle Struma Valley with the hottest place in Bulgaria,the sea coast, and a low mountain (Eastern Rhodopes). On the other hand, theselected regions include protected (Rila, Central Stara Planina) as well as unpro-tected (Belasitsa, Black Sea coast) territories, and the areas where limestone andkarst forms are present (Slavyanka, most part of Stara Planina) or absent (Rila,Struma Valley).
The main reasons for the selection of these regions are the following:Western and Central Stara Planina: their position on the dispersal route ofEurosiberian and Central European species to the Balkan Peninsula from theCarpathians; one of the two main routes of this dispersal;Rila: the best developed alpine belt, most strongly manifested mountain climate;Western Rhodopes: the best example of xerothermic and mountain fauna repre-sented in a mountain range;Belasitsa and Slavyanka: the only mountains in Bulgaria located as islands inthe Mediterranean Subregion;Middle Struma, Eastern Rhodopes, Black Sea coast: the warmest regions withrichly represented Mediterranean fauna.
The data on the species distribution were compiled using all available literaturefor Bulgaria. It includes major reviews (MR) as well as additional references (AR)
1 The official name of the park in English is Central Balkan National Park. In the present paper,the adjective “Balkan” is used only as an attribute for the Balkan Peninsula (Balkan Mediterranean,Montane Balkan, Balkan–Anatolian, Central Balkan etc.). Balkan is the Turkish name and StaraPlanina is the Bulgarian name of the mountain range dividing North and South Bulgaria. In orderto avoid confusion, for the mountain range we use only the name Stara Planina, e.g. Central StaraPlanina (Mts.), Eastern Stara Planina (Mts.), etc., and for the park, Central Stara Planina NationalPark.
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ORTHOPTERIDS OF BULGARIA 255
published earlier and later than these reviews, which list additional species fromthe respective region not mentioned in the major references. Below is the list ofthis literature.
Western Stara Planina. MR: Peschev (1970), Pešev (1974). AR: Buresch andPeschev (1957), Popov (1958), Peshev (1980, 1985), Andreeva (1985), Schmidt(1998).Central Stara Planina National Park. MR: Popov (2000a), Popov et al. (2000).AR: Pešev (1990), Popov (unpubl.).Rila National Park. MR: Popov (2000b).Western Rhodopes. MR: Pechev (1964, 1975). AR: Nedelkov (1908), Buresch(1939), Buresch and Peschev (1955, 1957, 1958), Bazyluk (1961), Popov (1970),Harz (1984, 1985b, 1986), Pešev (1990), Popov (1998).Eastern Rhodopes. MR: Popov and Chobanov (2004).Belasitsa. MR: Pešev (1962b). AR: Buresch and Peschev (1957), Pešev (1962a),Andreeva (1980), Peshev (1980), Peshev and Andreeva (1986), Popov (1997a),Popov (unpubl.).Slavyanka. MR: Pešev and Maran (1963). AR: Drenowski (1936, 1938), Bureschand Peschev (1957), Peshev (1980, 1981), Peshev and Andreeva (1986), Popov(unpubl.).Black Sea coast. MR: Peshev and Djingova (1974). AR: Drenowski (1938),Buresch (1939), Drensky (1942), Ramme (1951), Maran (1953), Buresch andPeschev (1957), Maran (1958), Bazyluk (1961), Popov (1970), Harz (1975),Karaman (1975), Andreeva (1985), Pešev (1990), Chobanov (2003), Popov(unpubl.).Middle Struma Valley. MR: Popov et al. (2001), only for Kresna Gorge.AR: Kaltenbach (1967), Harz (1975), Andreeva (1982), Popov and Ganev (1983),Harz (1985a, 1985b, 1986), Peshev and Andreeva (1986), Gorochov and Llorente(2001), Chobanov (2003).
The species reported in the major reviews, which occur next to but not insidethe certain region, were excluded after an analysis. The information on speciesdistribution is complete only for Kresna Gorge and the Eastern Rhodopes becausethe entire rich collection of the NMNHS from these regions was revised andidentified (Popov et al., 2001; Popov and Chobanov, 2004). The faunistic lists inthe oldest reviews, namely these for Belasitsa (Pešev, 1962b) and Slavyanka (Peševand Maran, 1963), are corrected. The species typical only for the lowlands but notoccurring in the mountain proper are deleted from these lists; the number of suchspecies is 34 for Belasitsa, and 20 for Slavyanka. We removed also five speciescharacteristic only for Strandja from the list of the Black Sea coast species byPeshev and Djingova (1974). The species added on the basis of publications otherthan the main reviews are most numerous for the Western Rhodopes (40 species),Middle Struma Valley (outside the Kresna Gorge) (34 species), and Black Sea coast(26 species).
Share of the fauna of the regions compared to the fauna of Bulgaria. Exactly50% of the Bulgarian taxa of Orthopterida occur in the Western Rhodopes. The
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256 A. POPOV
reason for such richness is the abovementioned combination of xerothermic andmountain fauna. In all other regions, less than 50% of the Bulgarian species arerecorded. Only the orders with the lowest species diversity are represented by alltheir Bulgarian species in some of the selected regions: Mantodea (four species)in Struma Valley and Eastern Rhodopes, Isoptera (two species) on the Black Seacoast, and Embioptera (one species) in Struma Valley and on the Black Sea coast.All three orders include only thermophilous species. Blattodea and Dermaptera arerepresented in each region at most by ca. 50% of their species. Their distributionin Bulgaria, however, is not yet well studied. Five small orders taken together havethe highest representation in the Struma Valley and the Black Sea coast (53% of theBulgarian taxa). Orthoptera has its highest representation in the Western Rhodopes(51% of the Bulgarian taxa).
Among the suborders, the Western Rhodopes are ranked first for the represen-tation of Ensifera (39% of the Bulgarian taxa), as well as Caelifera (71%). This richfauna of Orthoptera is due to the diverse habitats and the large area of the WesternRhodopes. The thermophilous and xerophilous fauna is concentrated in the valleysof the rivers in the northern part of the range (Chepinska Reka, Vacha, ChepelarskaReka), which flow from south to north. Before entering the Thracian Lowland andjoining the Maritsa River, these small rivers run through mountain gorges, whichprovide suitable refuges against the prevailing westerly winds for such species asTylopsis lilifolia, Sepiana sepium, Saga natoliae (Ensifera), Asiotmethis limbatus,Acrotylus patruelis, and A. longipes (Caelifera). The mountain fauna in the sameregion includes the inhabitants of the forest belts up to the coniferous belt. Thesubalpine zone occupies a limited area and is present only on Golyam PerelikPeak, where such mountain species occur as Isophya rhodopensis, Polysarcusdenticauda, Psorodonotus fieberi (Fig. 6), Anterastes serbicus (Ensifera), Podismapedestris, Omocestus viridulus, Myrmeleotettix maculatus (Fig. 2), Gompho-cerus sibiricus (Caelifera). The considerable difference in the share of WesternRhodopean Ensifera and Caelifera compared to all Bulgarian species is due to thebroader ecological plasticity of most Gomphocerinae, Oedipodinae, and Tetrigidae(Caelifera), which are widespread in Bulgaria, on one hand, and to the endemismand often sparse populations of many Phaneropteridae and Tettigoniidae (Ensifera),which are scantily represented in Bulgaria, on the other hand.
Among the families with high species richness, the same (and highest) share ofPhaneropteridae (30%) and Tettigoniidae (41%) compared to their total Bulgarianfauna is established in the Western Rhodopes and Western Stara Planina. Theexplanation for the species richness of Western Stara Planina is the same as forthe Western Rhodopes. There are refugia containing Submediterranean (expansiveMediterranean) fauna along Western Stara Planina such as the foot of this mountainrange near Belogradchik and the Iskar Gorge. Thermophilous species such asPhaneroptera nana, Leptophyes albovittata (Phaneropteridae), Platycleis affinis,Pholidoptera fallax (Tettigoniidae) occur there. The peaks of Sveti Nikolska Mts.and Berkovska Mts. (parts of Western Stara Planina) are inhabited by typicalmountain taxa such as Barbitistes serricauda (Fig. 2), Polysarcus denticauda,
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ORTHOPTERIDS OF BULGARIA 257
Poecilimon affinis affinis (Phaneropteridae), Metrioptera domogledi domogledi,M. helleri (Fig. 6), and Pholidoptera frivaldskyi (Tettigoniidae).
Among the subfamilies with high species richness, the Oedipodinae are bestrepresented in Struma Valley, and Gomphocerinae, in the Western Stara Planina bythe same share (76%) of their total Bulgarian fauna. The Struma Valley is inhabitedby both xerophilous (Celes variabilis, Sphingonotus caerulans, three species ofOedipoda, and three species of Acrotylus) and hygrophilous (Locusta migratoria,Aiolopus thalassinus, Mecostethus parapleurus, and Paracinema tricolor) Oedipo-dinae but all these species are thermophilous.
The lowest relative share of the Bulgarian fauna for the suborders Ensiferaand Caelifera is observed in Slavyanka with 21% and 35%, respectively. For thesubfamilies, this share is the lowest for Tettigoniinae in Belasitsa (27%), for Oedipo-dinae, in the Central Stara Planina National Park (24%), and for Gomphocerinae,in Slavyanka (31%). Slavyanka and Belasitsa, although closely located, are ratherdifferent in geological structure and vegetation. For that reason only 60% of thetotal number of orthopterid species found there inhabit both mountains. The lowrepresentation for the two mountains is due to their small area. The reasons for thelow species diversity of Oedipodinae in the Central Stara Planina National Parkare the lack of suitable habitats for hygrophilous and psammophilous species andthe relatively high altitude of large parts of this park.
Share of the main systematic groups compared to the entire orthopterid faunaof the regions. As main systematic groups here we consider the suborders, families,and subfamilies with high species richness. The share of Ensifera in the regionalorthopterid fauna is the lowest in the Rila National Park (37%). This is dueto the small share of Grylloidea and lack of the families Meconematidae andConocephalidae in the fauna of Rila. On one hand, only two most common speciesof Grylloidea, Gryllus campestris and Gryllotalpa gryllotalpa s.l., occur in thismountain range. On the other hand, Rila is the only region where Meconematidaeand Conocephalidae are not found. These facts can be explained by the high altitudeof the lower border of the park and mountain range, and by the absence of wetlandstypical for Conocephalidae, at the foothills of Rila. In other regions, Ensiferaconstitute 41–45% of all orthopterids.
Caelifera have the highest share in Western Stara Planina (54% of all orthopterids),due to the higher number of Gomphocerinae species (especially of the genusChorthippus) than in any other territory. Ch. montanus and Ch. vagans are knownin Bulgaria only from this region, but further verification of these data is necessary.
Among the families rich in species, the differences between the regions areinsignificant. The highest share of Phaneropteridae is observed in the Central StaraPlanina National Park (17%), and the lowest, in the Middle Struma Valley (9%).The share of Tettigoniidae is the highest in Slavyanka (22%), and the lowest, in theBlack Sea coast (16%).
More important are the differences in distribution of Oedipodinae and Gompho-cerinae. The poorest representation of Oedipodinae compared to all species is inthe national parks of Central Stara Planina (6%) and Rila (7%). The Struma Valley
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(14%) and the Black Sea coast and Belasitsa (13% each) are at the other extreme.Gomphocerinae have their minimal share in the Black Sea coast (15%), in theEastern Rhodopes (16%), and in the Struma Valley (17%); and the maximal share inthe national parks of Rila (33%) and Central Stara Planina (32%). The share of thesetwo subfamilies in the selected regions appears to depend not on the hygrophilyof the species but on their thermophily. The ratio of hygrophilous, mesophilous,and xerophilous species does not vary considerably among the regions. Accordingto their thermophily, however, these two subfamilies express the opposing trends.The thermophilous species prevail in Oedipodinae, and the cold-tolerant ones, inGomphocerinae. The regional distribution of species belonging to these two subfam-ilies is not surprising, keeping in mind the altitude of each region.
Species characteristic for certain regions. The most important characteristic of azoogeographical or biogeographical region is the complex of endemic (subendemic)species found only (or nearly only) in this region. In addition, it is important toanalyze other species, which for some reason (relict range, localities in the peripheryof their range, isolated records of very rare stenotopic species, etc.) are restrictedto a certain area within this territory. Below, we list the species characteristic fornine selected regions of Bulgaria; the species found only in the particular region arelisted under (A), and those found in this region and in one other region, under (B).1. Western Stara Planina.
(A): Metrioptera helleri (Fig. 6), Isophya miksici (both local endemics), andChorthippus montanus (the material on this species needs to be revised).
(B): Metrioptera domogledi domogledi (could occur only in this region), Isophyamodestior (also in Vitosha Mts.), Leptophyes discoidalis (also in KresnaGorge), Odontopodisma schmidti (also in Western Rhodopes), Poecilimonaffinis affinis, Pholidoptera littoralis, and Chorthippus vagans (the lastthree species with doubtful data for other regions).
2. Central Stara Planina National Park.(A): Isophya obtusa (a Balkan endemic), I. pravdini pravdini, and I. p. bazyluki
(an endemic species of Stara Planina with two local endemic subspecies inthis region).
(B): Eupholidoptera beybienkoi (currently considered a local endemic species;also in the Central Predbalkan; the taxonomic status of this species needsrevision).
3. Rila National Park.(A): Miramella sp. (the genus only in Rila as well), Barbitistes constrictus
(a relict population), Poecilimon affinis rilensis (an endemic subspeciesof Rila), Pholidoptera hoberlandti, and Ph. rhodopensis (both are localendemics; the taxonomic status of the last three taxa needs revision).
(B): Phyllodromica carniolica (Blattodea; also in Vitosha Mts.), Aeropedellusvariegatus (Fig. 1), and Isophya bureschi (both also in Pirin Mts.).
4. Western Rhodopes.(A): Isophya rhodopensis, Pholidoptera ganevi, and Stenobothrus bulgaricus
(all three are local endemics; the taxonomic status of the latter is unclear).
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ORTHOPTERIDS OF BULGARIA 259
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Aeropedellus variegatusBohemanella frigida
Fig. 1 Distribution in Bulgaria of the oreotundral species of Orthopterida: Bohemanella frigida andAeropedellus variegatus. The main geographical regions are named. Abbreviations: BEL, Belasitsa Mts.;MAL, Maleshevska Mts.; OGR, Ograzhden Mts.; OSO, Osogovo Mts.; SLA, Slavyanka Mts.; VIT,Vitosha Mts.; VLA, Vlahina Mts. Areas above 1200 m altitude are shaded.
(B): Odontopodisma schmidti (also in Western Stara Planina), Eumodicogryllusbordigalensis (also in Sofia), and Anechura bipunctata (Dermaptera; alsoin Rila Mts.; the last two species are placed in this category provisorily;their distribution in Bulgaria is insufficiently known).
5. Eastern Rhodopes.(A): Paranocarodes chopardi (a local endemic of the Bulgarian and Greek parts
of the mountain range; see Figs. 10 and 11), Duroniella laticornis (the onlylocalities of the species in Europe are in this region; see Fig. 4), Myrme-cophilus myrmecophilus, Xya pfaendleri, and Chorthippus porphyropterus(the last three species placed in this category provisorily; at least the lasttwo species are very likely widespread in Bulgaria but their distribution isnot studied).
(B): Iris oratoria (Mantodea) (Fig. 4), Eupholidoptera smyrnensis (both also inStruma Valley), Saga campbelli gracilis (also in Sakar Mts.; see Figs. 7and 8), Discoptila buresi (also in the Black Sea coast), and Chorthippusbornhalmi (also in Ograzhden Mts.; the last species placed in this categoryprovisorily; most likely widespread in Bulgaria but its distribution is notstudied).
6. Belasitsa.(A): Poecilimon kisi, P. belasicensis, P. mistshenkoi tinkae (all currently
believed to be local endemics), and Platycleis ebneri (a Balkan endemic).
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260 A. POPOV
(B): Pholidoptera macedonica (also in Slavyanka Mts.) and Saga campbellicampbelli (also in Struma Valley with the adjacent part of Slavyanka Mts.;see Fig. 8).
7. Slavyanka.(A): Poecilimon mistshenkoi marzani (a local endemic subspecies).(B): Isophya kisi (also in Pirin Mts.), Gampsocleis abbreviata (also in
Sakar–Tundja Region), Saga campbelli campbelli (Fig. 8), and Pholi-doptera macedonica (about the last two taxa see above under Belasitsa).
8. Black Sea coast.(A): Anisolabis maritima (Dermaptera), Kalotermes flavicollis (Isoptera)
(Fig. 4), Epacromius coerulipes, Platypygius crassus, Dociostaurus kraussi,and (introduced) Meconema meridionale.
(B): Haploembia solieri (Embioptera) (Fig. 4), Epacromius tergestinus,Modicogryllus truncatus (also in Struma Valley; the distribution of thelatter in Bulgaria is insufficiently known), Isophya modesta (also in theadjacent parts of Dobrudja; see Fig. 6), Discoptila buresi (also in the EasternRhodopes), Paranocarodes straubei (also in the adjacent part of StrandjaMts.; an isolated population near Sliven has an uncertain taxonomic status;see Fig. 11), and Forficula smyrnensis (Dermaptera; also in the adjacentparts of Strandja Mts. and in Ograzhden Mts.).
9. Struma Valley.(A): Apterygida media (Dermaptera), Polyphaga aegyptiaca (Blattodea),
Isophya andreevae (a local endemic), Platycleis macedonica (a Balkanendemic), Andreiniimon nuptialis, Pteronemobius heydeni tartarus,Pallasiella turcomana, Eupholidoptera marani, Tartarogryllus sandanski(two last species are local endemics; their taxonomic status needs revision),and Dociostaurus tartarus (the material on this species needs to be revised).
(B): Dociostaurus genei (Fig. 4), Gryllomorpha cf. miramae (both species alsowith doubtful data for the Rhodopes; perhaps occur only in the StrumaValley), Leptophyes discoidalis (also in Western Stara Planina), Iris oratoria(Mantodea) (Fig. 4), Eupholidoptera smyrnensis (both also in the EasternRhodopes), Saga rammei (also near Popovitsa in the Thracian Lowland, seeFig. 8), Haploembia solieri (Embioptera) (Fig. 4), Epacromius tergestinus,Modicogryllus truncatus (about the last three taxa see above under Black Seacoast), and Saga campbelli campbelli (see above under Belasitsa, Fig. 8).
3·2 Vertical distribution
Vertical distribution of species is described along the altitudinal vegetationbelts, which are commonly accepted by the botanists for Bulgaria. Accordingto the localities and ecological requirements of every species, it is estimated towhich of the belts each record belongs. The vegetation belts used here are thefollowing: xerothermic oak forests: Quercetum; mesophilous oak and mixed forests:
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Quercetum, Carpinetum; beech forests: Fagetum; coniferous forests: Pinetum (Pinussylvestris), Piceetum; subalpine vegetation: Pinetum (Pinus mugo), Juniperetum;and alpine vegetation: Caricetum, Seslerietum. The total number of taxa recordedfor each belt is shown in Table 2. Not surprisingly, these numbers decrease fromthe lowest to the highest altitudinal belt.
All orders, suborders, superfamilies, families, and subfamilies of Orthopteridarecorded for Bulgaria are found in the xerothermic oak belt (Table 1). Only 13 outof 112 Bulgarian orthopterid genera are absent from that lowest belt. One of thesegenera, Apterygida, belongs to Dermaptera and the rest to Orthoptera. Twelve ofthese genera are represented in the Bulgarian fauna with one species each, and onlyBarbitistes, with two species. The xerothermic belt is inhabited by 202 species andsubspecies (over 75% of the Bulgarian taxa). In every other belt, less than 50%of the Bulgarian taxa are found (Table 2). This richness is due to a considerableshare of southern thermophilous elements in the Bulgarian orthopterid fauna, whichare found only in the lowlands, and often only in South Bulgaria or even only inits southernmost and warmest parts. This trend is illustrated by a strong decreaseof the species number per area unit in the direction Greece–Bulgaria–Romania–Hungary–Slovakia–Poland. Many examples of such species can be easily found inTable 1.
Only in the xerothermic belt occur the following taxa: the order Embioptera;the families Polyphagidae, Kalotermitidae, Mogoplistidae, Myrmecophilidae, andAnisolabididae; the subfamilies Copiphorinae, Onconotinae, Gryllomorphinae,Akicerinae, and Cyrtacanthacridinae (Table 1); and 81 species and subspecies(Table 3). The abovementioned order, families, and subfamilies, however, arerepresented in Bulgaria only by one species each, except Gryllomorphinae (threespecies). Among the species-rich superfamilies of Orthoptera, the share of taxainhabiting only the xerothermic belt as compared to all Bulgarian taxa varies from27% (Tettigonioidea and Acridoidea) to 68% (Grylloidea).
The number of taxa in the mesophilous oak belt and the beech belt is quite close(Table 2). However, only 66% of the taxa present in the beech belt are found alsoin the mesophilous oak belt (78 of 119 taxa). Among the other pairs of adjoiningbelts, this share varies from 79% for the subalpine and coniferous belts to 86%
Table 2 Vertical distribution of Orthopterida in Bulgaria by vegetation belts
Vegetation belt Number of taxa Share from all Bulgariantaxa (%)
xerothermic oak forests 202 76mesophilous oak forests 124 46beech forests 119 45coniferous forests 82 31subalpine vegetation 43 16alpine vegetation 10 4
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Table 3 Upper limit of vertical distribution of Orthopterida in Bulgaria by vegetation belts
Vegetation belt Number of taxa Share from all Bulgariantaxa (%)
xerothermic oak forests 81 30mesophilous oak forests 46 17beech forests 49 18coniferous forests 48 18subalpine vegetation 33 12alpine vegetation 10 4
for the mesophilous and xerothermic oak belts and even 100% for the alpine andsubalpine belts, i.e. all species of the alpine belt occur also in the subalpine belt.From another perspective, the taxa found in the beech belt constitute 45% of allBulgarian orthopterids, and the species common between the beech belt and themesophilous oak belt constitute only 29% of Bulgarian fauna. For other pairs ofadjoining belts, the difference between these shares is only 3–6%. Therefore, theboundary between the mesophilous oak belt and the beech belt has to be accepted asan important barrier for the dispersal of the orthopterid fauna. This boundary runs at900–1000 m a.s.l. in the major part of Bulgarian mountains except the southernmostones. Below this boundary one finds the warm and dry oak forest; above it, thecold and humid beech forest.
The maximum similarity of the faunistic diversity (number of common species)is found in the xerothermic and mesophilous oak belts as well in the beech andconiferous belts. In the first case, there are 107 common taxa (86% of 124 taxafound in the mesophilous oak belt). In the second case, there are 69 common taxa(84% of 82 species found in the coniferous belt). This similarity is due to therather similar conditions in temperature and humidity in the xerothermic oak andmesophilous oak belts. The conditions in the beech and coniferous belts are alsosimilar, although very different from those in the oak belts. Table 3 shows the closenumber (46 to 49) of species and subspecies with the upper limit of distribution inthe mesophilous oak, beech, and coniferous belts.
The comparison of the number of taxa found only in a single vegetation belt(Table 4) indicates a very high number of species and subspecies characteristicfor the xerothermic oak belt. This can be explained by the rich representation ofMediterranean orthopterid species in Bulgaria. The higher specificity of taxa in thebeech belt compared to the other belts is due to the wide distribution of the beechforests in Stara Planina, Sredna Gora, and in the smaller Bulgarian mountains. Atthe same time, the subalpine and alpine belts are limited in area, and a developedconiferous belt exists only in about the half of the high mountain ranges in Bulgaria(Rila, Pirin, Western Rhodopes, Vitosha, and Slavyanka).
The timberline (the boundary between the coniferous and subalpine belts), alsorepresents an important border for the species distribution. It runs in the Bulgarian
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Table 4 Taxa of Orthopterida in Bulgaria, found only in one vegetationbelt (stenotopic in vertical distribution)
Vegetation belt Number of taxa
xerothermic oak forests 81mesophilous oak forests 6beech forests 12coniferous forests 7subalpine vegetation 6alpine vegetation −
mountains usually at the altitude of 1800–2000 m a.s.l.; rarely, single coniferoustrees reach 2200 m. The subalpine belt is developed in the eight highest mountainranges in Bulgaria: Rila, Pirin, Stara Planina (in its western and central parts),Vitosha, Osogovo, Belasitsa, Slavyanka, and Western Rhodopes (only around thepeaks of Golyam Perelik and Syutkya). The alpine belt is found only in Rila andPirin above 2500 m and around Botev Peak in Central Stara Planina. Eight speciesof Bulgarian orthopterids are found entirely above the timberline. Two of themoccur in the subalpine and alpine belts: these are the only species with Arctoalpinedistribution and representatives of oreotundral fauna among Bulgarian Orthoptera,Bohemanella frigida and Aeropedellus variegatus (Fig. 1). The remaining specieshave been found only in the subalpine belt. The timberline in Belasitsa is situatedon 1700 m; Poecilimon mistshenkoi tinkae, P. kisi, and P. belasicensis occur fromthis altitude up to 1900 m. P. pechevi is found in the treeless high mountain zoneof Vlahina Mts. above 1600 m, and Metrioptera helleri, above the timberline onTodorini Kukli Peak in Western Stara Planina at 1640 m (Fig. 6). All these taxa,characteristic only for the subalpine belt, are local endemics. Isophya brevipennis isrecorded in the subalpine belt of Pirin (Köhler, 1988) but its occurrence in Bulgarianeeds confirmation since this record is based only on a female.
All species inhabiting the alpine belt occur also in the subalpine belt. Eightof the species, distributed together above and below the timberline, occur up tothe alpine belt, and 17 taxa occur in the subalpine belt and below, many of themin all belts. These are eurytopic species with broad ecological plasticity. Typicalrepresentatives reaching the alpine belt are Stenobothrus nigromaculatus (from600 m up to 2600 m) and Chorthippus brunneus (0–2600 m) (the group of speciesof the latter needs revision). More species occur in all belts except the alpineone; typical examples are Isophya speciosa, Poecilimon thoracicus, Omocestushaemorrhoidalis, and Chorthippus parallelus, distributed from the seashore up to2300–2500 m.
The maximum altitude of occurrence of the orthopterid orders in Bulgaria isshown in Table 5. For the orders found in Bulgaria also in the mountains, themaximum altitude is recorded in the south of the country (Pirin Mts. and SlavyankaMts). These two mountain ranges are formed of limestone and marble, and therefore
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Table 5 Highest altitudinal occurrence of the orthopterid orders in Bulgaria
Order Highest altitude (m) Region Species
Orthoptera 2850 Pirin Anterastes serbicusOrthoptera 2850 Pirin Bohemanella frigidaOrthoptera 2850 Pirin Gomphocerus sibiricusOrthoptera 2850 (2900) Pirin (Rila) Aeropedellus variegatusBlattodea 2000 Rila, Pirin, Slavyanka Ectobius balcaniMantodea 1650 Slavyanka Mantis religiosaDermaptera 1600 Slavyanka Forficula auriculariaIsoptera 1000 Stara Planina Reticulitermes lucifugusEmbioptera 200 Kresna Gorge Haploembia solieri
characterized with higher temperatures. Isoptera and Embioptera, both practicallyentirely tropical groups, also reach high in Bulgaria in sunny and warm localities.
4 Zoogeography
Zoogeographical identity of the Bulgarian orthopterid taxa is discussed here in twoaspects: the type of distributional range (chorotype) and the suggested category oforigin (i.e. center of dispersal, speciation and possible origin). Placement of eachspecies and subspecies in a certain category is a result of the original analysis ofexisting ranges. In many cases, categories proposed previously for zoogeographicalcharacterization of Orthopterida or other animal groups are not used and an originalscheme is provided.
4·1 Chorotypes
The species are grouped according to their distribution into a detailed system ofchorotypes; only the most similar or identical ranges fall under the same chorotypecategory. Although it is better to use a smaller number of categories to classifygeographical ranges, the high degree of endemism in Orthoptera, and the presenceof other species with restricted ranges, necessitate introduction of a large numberof categories in order to interpret and compare the species distribution in detail.The chorotypes are arranged in groups according to the size of their ranges. Thecategories proposed here and their content were not previously used in the zoogeo-graphical literature.
Lately, the chorotypes of Vigna Taglianti et al. (1999) for the WesternPalearctic have been increasingly used in the Bulgarian zoological literature.However, their scheme does not correspond well to the typification of theranges either for the species in the Balkan Peninsula or of those in WesternPalearctic. Many examples can be given but the following two are sufficient to
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support this statement. The typical species with Eurosiberian distribution occurin the east to Manchuria (Tettigonia cantans, Arcyptera microptera, Gompho-cerus sibiricus, Gomphocerippus rufus, Chorthippus apricarius, Ch. montanus,and Ch. dorsatus), Korea (Psophus stridulus and Omocestus haemorrhoidalis),Primorye Region (Chrysochraon dispar), the islands of Sakhalin and Kunashir(Chorthippus biguttulus), and Japan (Phaneroptera falcata and Mecostethusparapleurus). In the scheme of Vigna Taglianti et al. (1999), however, such speciesdo not correspond to the Siberian–European chorotype but instead belong to thecategory of Asian–European species. The other example concerns the species withCarpathian–Balkan distribution (Pholidoptera frivaldskyi), Southeastern European(Odontopodisma decipiens, Platypygius crassus, and Omocestus minutus), andSoutheastern European–Western Asian distribution (Onconotus servillei, Pallasiellaturcomana, Dociostaurus brevicollis, and D. kraussi). According to Vigna Tagliantiet al. (1999), they should belong to the Turanian–European chorotype. All thesespecies, however, do not occur in the Turanian Subregion or in the SouthwesternEurope. Some of them are steppe species such as Onconotus servillei (Fig. 3),Dociostaurus brevicollis, D. kraussi, and the halophilous Platypygius crassus, andtheir placement in the Turanian–European chorotype is incorrect. It should bementioned also that the classification of Vigna Taglianti et al. (1999) uses manychorotype names which do not seem to be correctly formed in English.
To determine the chorotypes of Bulgarian endemic species and subspecies,the categories of Hubenov (1997) for division of Bulgaria were often used. Thescheme of Dr. Zdravko Hubenov, developed in order to designate the distributionof the taxa in Bulgaria, is based on the physical-geographical structures. Theboundaries of the natural geographical territorial units are outlined in this schemevery precisely but the level of subdivision does not always correspond to thesimilarity and difference in the species diversity. On one hand, areas similar intheir faunas are separated at the first level (as regions), e.g. Central Danubian Plainand Central Predbalkan, which have similar fauna and the same endemic speciesIsophya plevnensis, or the Black Sea coast and Dobrudja that share the endemicsubspecies I. modesta longicaudata (Fig. 6). On the other hand, areas with verydifferent faunas are separated only at the third level (as districts), e.g. WesternRhodopes and Eastern Rhodopes as well as Osogovo Mts. and Middle StrumaValley.
We grouped the Bulgarian species of Orthopterida into 49 chorological categories(chorotypes) (Table 6). In reality, the number of chorotypes could be smaller. Themore complete faunistic exploration of Bulgaria and adjacent countries could lead tothe removal of some regions that will be left without a single characteristic Bulgarianendemic taxon, e.g. some mountain ranges (Vlahina, Maleshevska, Ograzhden,mountains around Sofia, Eastern Stara Planina) and of the provisional regions withcomplicated structure and compound names (Eastern Rhodopean–Black Sea coast,Macedonian–Rhodopean–Black or Marmara Sea coast, northern part of BulgarianBlack Sea coast). In addition, some taxa could be found to be synonyms of speciesand subspecies from Bulgaria or adjacent countries. As a result, the number of
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Table 6 Chorotypes of Orthopterida in Bulgaria
Chorotypes Number of taxa Share from allBulgarian taxa (%)
Holarctic Oreotundral 2 0�7Cosmopolitan 10 3�7Holarctic 1 0�4Palearctic 9 3�3Western Palearctic 5 1�86Eurosiberian 33 12�2European–Western Asian 8 3�0European 6 2�2Central European 3 1�1Central and South European 29 10�7South European 5 1�86Southeastern European 6 2�2Southeastern European–Western Asian 7 2�6Holomediterranean 15 5�6Transadriatic 4 1�5Northwestern Balkan 4 1�5Northern and Central Balkan 5 1�86Central Balkan 4 1�5North Bulgarian 1 0�4Central Danubian–Central Predbalkan 1 0�4northern part of Bulgarian Black Sea Coast 2 0�7Western Stara Planina 2 0�7Central Stara Planina 3 1�1Eastern Stara Planina 1 0�4Sofia Mountainous 1 0�4Central Sredna Gora 1 0�4Osogovo 1 0�4Vlahina–Maleshevska–Ograzhden 3 1�1Belasitsa 3 1�1Rila–Pirin–Slavyanka 9 3�3Western Rhodopean 3 1�1Eastern Rhodopean 1 0�4Eastern Rhodopean–Black Sea Coast 1 0�4Thracian–Black Sea Coast 2 0�7Tundja 1 0�4Strandja 2 0�7Middle Struma Valley 3 1�1Eastern Balkan 5 1�86Macedonian 4 1�5Macedonian–Rila–Rhodopean 2 0�7Macedonian–Rhodopean–Black or Marmara Sea Coast 2 0�7Macedonian–Thracian 1 0�4Southern Balkan 5 1�86Balkan–Anatolian 13 4�8Eastern Mediterranean 16 5�9
continued
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Table 6 continued
Chorotypes Number of taxa Share from all Bulgariantaxa (%)
Turanian–Mediterranean 11 4�1Turanian–Caucasian 1 0�4Afrotropical–Palearctic 9 3�3Paleotropical–Palearctic 4 1�5
chorotypes as well as the number of taxa could be reduced. Furthermore, one of thecategories in the list is represented by a sole species, which is very likely incorrectlyidentified.
Most numerous are the taxa with Eurosiberian distribution (Table 6), although thethermophilous and southern species in general prevail among the orthopterid insectsin Bulgaria. That is due to the fact that the Eurosiberian species form a homogeneousgroup with similar ranges while the Mediterranean species are scattered among alarge number of chorotypes. Ranked by the species number, the next five placesafter the Eurosiberian taxa are taken by chorotypes with thermophilous fauna (fromCentral and South European to Turanian–Mediterranean).
The ranges of 244 taxa, which are classified in 44 chorotypes, fall entirelywithin the borders of the Palearctic. The remaining 26 species recorded in Bulgarianow or in the past belong to five chorotypes. There are three Holarctic orHolarctic Oreotundral species whose ranges do not exceed the borders of theHolarctic. The species of Cosmopolitan, Paleotropical–Palearctic, and Afrotropical–Palearctic chorotypes are represented in Bulgaria by a total of 23 species; they aredistributed in the Holarctic and beyond its borders.
The chorotypes listed in Table 6 can be arranged in three groups accordingto the range size. The group with wide ranges includes chorotypes reaching inthe east to the Far East or Siberia, as well as those exceeding the borders of thePalearctic. The group with medium sized ranges unifies chorotypes covering theentire Europe, or Central and South Europe, and adjacent regions in the east tothe Western or Central Asia. The group with small ranges consists of chorotypesincluding only South Europe or parts of it and Anatolia. The first group ofwidespread species includes only seven chorotypes (from the Cosmopolitan to theEurosiberian) with 68 species (25.2% of all Bulgarian species). The other twogroups include the same number of taxa (each has 101 species and subspecies,or 37.4% of the Bulgarian fauna) but a higher number of chorotypes. The groupwith medium-sized ranges consists of 10 chorotypes (from the Western Palearcticto the Central and South European), and the group with small-sized ranges unifies32 chorotypes (65% of all 49 chorotypes in the list). This division reflects the factthat the species with restricted distribution predominate in the Bulgarian orthopteridfauna.
The Cosmopolitan chorotype includes three groups of species. Three species ofBlattodea and Acheta domesticus are synanthropic. Four species of Dermaptera are
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also introduced accidentally, mostly by cargo boats, to all parts of the world. Dueto their ecological plasticity, they have adapted to a great variety of conditions inall or nearly all continents. Only Aiolopus thalassinus has a Cosmopolitan (in fact,Subcosmopolitan) range, which is not a result of an anthropogenic introduction. Itoccurs almost everywhere in the East Hemisphere except the northernmost regions.
Only Tetrix subulata has a Holarctic distribution. The almost entire absence ofHolarctic species among the Orthopterida in Bulgaria is impressive when comparedto other animal groups.
The distinction between the distribution of Palearctic and Eurosiberian speciesis rather small, particularly in the Asian part of their ranges.
The European part of the range of the European–Western Asian chorotypeincludes the entire Europe, e.g. Conocephalus dorsalis and Sphingonotus caerulans,or only Central and South Europe, e.g. Gampsocleis glabra and Saga pedo.
Species found only in Europe, e.g. Apterygida media (Dermaptera) andMeconema thalassinum, as well as species found in Europe and Anatolia, e.g. Pholi-doptera griseoaptera, are interpreted here as European. The land bridge betweenthe Balkan Peninsula and Anatolia was severed only ca. 10,000 years ago, and themodern sea straits are not a true barrier preventing the mixing of the two faunas.Thus, there is no principal difference between species occurring in Europe plusAnatolia, or only in Europe.
As Transadriatic, we characterize the taxa found only in the Apennine partof Italy (the Apennine Peninsula proper, without North Italy) and in the BalkanPeninsula, e.g. Andreiniimon nuptialis and Eupholidoptera chabrieri schmidti.The species occurring on the Apennine Peninsula, Balkan Peninsula, and inAnatolia, e.g. Platycleis nigrosignata, are treated here not as Transadriatic but asEastern Mediterranean together with the species inhabiting the Balkan Peninsulaand the eastern parts of the Mediterranean, e.g. Ameles heldreichi (Mantodea),Saga natoliae, and Duroniella laticornis. The species found only in the BalkanPeninsula and Anatolia, e.g. Phyllodromica pallida (Blattodea), Eupholidopterasmyrnensis, and Bucephaloptera bucephala, are separated as the Balkan–Anatolianchorotype.
The only Turanian–Caucasian species is Dociostaurus tartarus. A revision ofthe material will most likely show that this species does not occur in Bulgaria, andthis chorotype category will have to be deleted.
The Afrotropical–Palearctic and Paleotropical–Palearctic species include not onlyspecies originating from the tropical areas of the Old World. These categoriesinclude, respectively, the Holomediterranean by origin Anacridium aegyptium andOedaleus decorus which ranges to the Afrotropical Region, and species of the genusXya, for which the distribution in the Paleotropical regions is insufficiently knowndue to the taxonomic problems.
There are nine Balkan chorotypes (excluding those confined entirely in theterritory of Bulgaria), with 32 species and subspecies (Table 6), evenly distributedamong these chorotypes. If we pool the three chorotypes derived from theMacedonian chorotype in one, the seven resulting categories will include 4 to 5
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taxa each. Nearly all these taxa are Balkan endemic species and subspecies. Theranges of the remaining taxa extend only slightly beyond the borders of the BalkanPeninsula.
There are 19 Bulgarian chorotypes with 41 species and subspecies (Table 6).Most of these taxa are Bulgarian endemic species and subspecies; only a few ofthem have ranges slightly extending beyond the territory of Bulgaria. The Rila–Pirin–Slavyanka chorotype is the most diverse, with nine endemic species andsubspecies (at least three times more than any of other Bulgarian chorotypes).
4·2 Origin and dispersal
To interpret the place of species’ origin, one considers not only the recent rangesbut also the suggested place of origin of the genera (and for the subspecies, alsothe origin of the species). This analysis is initially based on the theory of zoogeo-graphical categories proposed for the Holarctic by de Lattin (1967), in whicheach species can be treated as a faunal element within three biochores (high-levelbiomes): Arboreal, Eremial, and Oreotundral. The types of zoogeographical centersconsidered by de Lattin (1967) are: center of (ancestral) origin, center of differen-tiation, center of preservation (refugium center), and center of dispersal. Determi-nation of the centers of origin and differentiation on the basis of distribution of acertain systematic group is quite difficult and uncertain. However, determination ofthe centers of preservation and dispersal could be more feasible since it is associatedwith the latest dynamic events (expansion or contraction), which resulted in therecent range. In some cases, the ranges have not undergone considerable changes,and are limited to the respective second-level (e.g. for the Pontomediterraneanstationary species) or third-level (e.g. for the Balkan species) center of dispersal;or the route of the expansion is commonly accepted, e.g. for the Siberian species.In many cases, only the first-level center of speciation and dispersal, e.g. for theHolomediterranean species, is known.
Four of the categories of origin according to de Lattin (1967) are used here:Siberian, Mongolian, Mediterranean, and Caspian faunal elements. The content ofthe last category, however, is changed since we include here the steppe species withdistinct type of range. Most remaining categories of origin are proposed here assubdivisions of the Mediterranean center of dispersal and of its Pontomediterraneanportion. Not only the ranges but the habitats and ecological requirements are takeninto consideration in this analysis. There are 24 categories of origin; 21 of theminclude only autochthonous (non-introduced) correctly identified species.
Eremial fauna. There are no representatives of the eremial fauna among theBulgarian orthopterids. Acrotylus longipes takes a transitional position betweenthe arboreal and eremial fauna. It is found in the Adriatomediterranean andPontomediterranean arboreal centers and in almost the entire Afrotropical Regionsouthward to Zambia and Namibia. The Bulgarian name of the species is DuneGrasshopper. This psammophilous species is adapted to the desert and semidesert
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areas in Africa as well as arid sandy and gravel seashore and river banks withoutvegetation in Bulgaria. The other two species of the same genus in Bulgaria,A. insubricus and A. patruelis, occur also in the entire Africa, including Sahara andNamibia, and in the desert areas of Arabia and Central Asia. They also inhabit drysandy and rocky terrains with very scanty vegetation but according to their habitatsin Bulgaria they are closer to arboreal species than A. longipes. Both species areamong the few in Europe that hibernate as adults, which is also a consequence oftheir tropical origin.
Oreotundral fauna. Oreotundral fauna by origin does not exist. Oreotundral(Arctoalpine) distribution is characteristic for some strongly expansive animalspecies with tundral or oreal origin. Only two species, Bohemanella frigida andAeropedellus variegatus, represent this fauna among the Orthoptera in Bulgaria(Fig. 1). The northern, continuous part of their Arctoalpine ranges prevails stronglyover the southern mountain part and covers the entire extreme North of Palearctic(B. frigida) or its Siberian part and Finland (Ae. variegatus). The southern mountainpart of the ranges includes the Alps, the mountains of Bulgaria, and Altai Mts. (bothspecies), some other Central European and Balkan mountains and Southern Siberia(first species), the Apennines and Caucasus (second species). In the Nearctic, thesetwo species occur only in Alaska (with the former also in Northwestern Canada).This indicates at least for Aeropedellus variegatus that its dispersal from East Asiavia Bering landbridge has stopped at its initial stage. The chorotype of both specieshas to be identified as Holarctic Oreotundral and the origin at least of Ae. varie-gatus, as Siberian Tundral. The migration of Bohemanella frigida was realizedmost likely in the opposite direction (see below under Distribution and origin ofselected genera). Several subspecies of these two species have been described (forB. frigida see also below under Distribution and origin of selected genera). TheBulgarian populations of B. frigida (Fig. 1) correspond to the subspecies B. f. strandi(Fruhstorfer, 1921) described from the Alps. For most Arctoalpine species, thetime since separation of the northern and southern populations is not consideredto be sufficient for differentiation of subspecies. Thus the question whether somepopulations represent distinct subspecies is left open.
Among other inhabitants of the high mountains, six species and subspecies occuronly in the subalpine belt, and eight other species reach up to the alpine belt (seeabove under Vertical distribution). All these taxa, however, are representatives ofarboreal but not of oreal fauna.
Arboreal fauna. Almost all (99.3%) of the Bulgarian orthopterid species belongto the arboreal fauna. Thamnobionts (dendrobionts) are only a small part of thesetaxa. Considerably more species are chortobionts, which inhabit clearings in themountain forests but also meadows and other open areas as well as arid, treelesshabitats. These open areas are secondary deforested habitats or primary steppes.Because of that, and judging from their general distribution, these chortobionts arealso typical arboreal species by origin.
The origin remains unclear for Modicogryllus truncatus and for both species ofTridactyloidea due to taxonomic problems and lack of revision of distributional data,
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as well as for two species of Dermaptera because of their anthropogenic dispersalnearly all over the world. The category of origin is assigned to the remaining 265species, but for 16 of them with some reservations. The species with uncertainlyinterpreted origin belong to Blattodea (one species), Tettigonioidea (seven species,including both species of Bradyporinae), Grylloidea (two species), and Acridoidea(six species), identified as Caspian, Pontomediterranean, Dinaric, Montane Balkan,Moesian, Thracian, and Anatolian.
According to their origin, the arboreal species of Orthopterida in Bulgariaare divided into 23 categories (Table 7). Three of these categories, namelyAdriatomediterranean, Iranian, and Sinotibetan faunal elements, are not typicalfor the Bulgarian fauna and include introduced or most likely incorrectly
Table 7 Zoogeographical origin of Orthopterida in Bulgaria
Categories of origin Number of taxa Share from all Bulgariantaxa (%)
OREOTUNDRALSiberian Tundral 2 0�8
ARBOREALNorthern arboreal
Siberian 35 13�2Siberian–Nearctic 1 0�4Siberian–Mediterranean 8 3�0Mongolian 1 0�4Central European 27 10�2Central European–Mediterranean 3 1�1Caspian 12 4�5
BalkanDinaric 9 3�4Montane Balkan 9 3�4Moesian 6 2�3Stara Planina 10 3�8Rila–Rhodopean 21 7�9Macedonian 11 4�1Thracian 11 4�1Balkan Mediterranean 7 2�6
MediterraneanHolomediterranean 37 14�0Pontomediterranean 30 11�3Anatolian 8 3�0
TropicalAfrotropical 7 2�6Paleotropical 6 2�3
Species introduced or unconfirmed in BulgariaAdriatomediterranean 2 0�8Iranian 1 0�4Sinotibetan 1 0�4
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identified species. The richest in species categories are Holomediterranean, Siberian,Pontomediterranean, and Central European elements (Table 7), which include 49%of all Bulgarian species.
The arboreal faunal elements in Bulgaria (Table 7) can be arranged in threegroups according to the position of their centers of dispersal towards the territory ofBulgaria. The species of the categories originating from the Siberian, Mongolian,Central European, and Caspian centers of dispersal are considered to be of northernorigin. The species originating from the Mediterranean and tropical areas areconsidered to be of southern origin. The species with Balkan origin are placedseparately in Table 7. These three groups are almost equal in species number:northern, 87; southern, 88; and Balkan, 84 (32 to 33% of all Bulgarian taxa in eachgroup). These categories do not include two oreotundral species of northern originand four introduced and unconfirmed species of southern origin.
The question arises about the place of the taxa with Balkan origin. Three outof eight Balkan centers of speciation (Macedonian, Thracian, and Balkan Mediter-ranean) are located in the Mediterranean Subregion (Fig. 5), i.e. they fall underthe categories with southern origin. Even among these faunas, however, there arethree taxa with Macedonian origin which in fact also inhabit localities outsideof Mediterranean Subregion. Pholidoptera macedonica occurs in Slavyanka above900 m up to the highest peak of this mountain. Saga campbelli campbelli (Fig. 8)and Poecilimon zwicki (Fig. 6) are also found in Slavyanka up to 1600 and 1500 m,respectively. Other five Balkan centers of speciation are located entirely in theEurosiberian Subregion and their faunas should be considered to have northernorigin. In the theory of zoogeography, all Balkan and Bulgarian endemic taxawere earlier considered a special case of Pontomediterranean faunal elements(Gruev, 1988). Later, the same author revised his concept treating (in our opinion,correctly) the endemic taxa as faunal elements different in origin and belongingto different categories (Gruev and Kuzmanov, 1994). Here, the categories of taxaoriginating from the Balkan centers of speciation are separated in a special group.The great majority of these taxa is closely related to southern species and, inaddition, includes representatives of genera with southern origin, e.g. Ancistrura,Andreiniimon, Gampsocleis, Saga, Discoptila, and Paranocarodes. Few are thetaxa closely related to northern species, e.g. representatives of Metrioptera and ofsubfamily Catantopinae. In the genera Isophya and Poecilimon, an intensive speci-ation in the extra-Mediterranean areas (e.g. Bulgaria) of the ancestral taxa, whichdispersed from the Mediterranean areas, has taken place. All this indicates that mostof the Balkan taxa have to be treated as taxa of southern origin. Therefore, specieswith southern origin in Bulgaria prevail to a large degree over those with northernorigin.
Division according to the origin differs among particular systematic groups. Allspecies of Mantodea, Isoptera, and Embioptera in Bulgaria have southern origin.All these orders have tropical origin from which only a small number of represen-tatives reaches South Europe. Southern and Balkan taxa predominate strongly inTettigonioidea, while northern species predominate in Caelifera. The share of the
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ORTHOPTERIDS OF BULGARIA 273
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Fig. 2 Distribution in Bulgaria of some species of Orthoptera with northern arboreal origin. Siberianfaunal elements: Phaneroptera falcata, Chrysochraon dispar, Euthystira brachyptera, Myrmeleotettixmaculatus. Central European faunal elements: Barbitistes serricauda, Pholidoptera griseoaptera,Odontopodisma rubripes, Stenobothrus crassipes. Areas above 1200 m altitude are shaded.
taxa with northern origin compared to all Bulgarian species of each group is 22%in Tettigonioidea, and 55% in Caelifera.
The richest in species in the group of northern arboreal species are the Siberianones (40%), followed by the Central European and Caspian faunal elements(Table 7). The Siberian elements (Fig. 2) are grouped in one category becausetheir origin from Ussurian, Sinokorean, or Angarian subcenters of origin cannotbe distinguished. They all are believed to be young species, which additionallydispersed recently from the Siberian center. Examples of species with rangesreaching eastward to Manchuria, Korea, Primorye Region, Sakhalin, and Japanare given above under the Chorotype section. The ranges of many other speciesdo not reach the Pacific Ocean in the east, e.g. Ectobius lapponicus (Blattodea),Conocephalus dorsalis, Arcyptera fusca, and Omocestus viridulus. There arethermophilous Siberian elements among the orthopterids, e.g. Anechura bipunctata(Dermaptera), Platycleis veyseli, and Mecostethus parapleurus. Their ranges arelocated more southward from those of the typical Siberian species and they often donot reach the East Siberia. Gomphocerus sibiricus is a Siberian faunal element withBoreomontane distribution. Its range is disjunct between the lowland of North and
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Central Russia, Siberia, and Kamchatka and the mountains of South Europe andCentral Asia. Disjunction in this case is smaller than in the Arctoalpine ranges. TheSiberian species have dispersed to the Balkan Peninsula via Dinaric Alps and viaCarpathians. More cold-tolerant species are found in Bulgaria only in the mountains,e.g. Tettigonia cantans, Metrioptera bicolor, Myrmeleotettix maculatus (Fig. 2), andOmocestus viridulus. More ecologically plastic species inhabit both lowlands andmountains, e.g. Omocestus haemorrhoidalis, Stenobothrus lineatus, S. nigromac-ulatus, and Chorthippus parallelus, all of them from subfamily Gomphocerinae.Most thermophilous Siberian species are found in Bulgaria only in the lowlands.
The Siberian–Mediterranean species are assumed here to have an initial and anadditional center of dispersal. In this scenario, they have dispersed to Europe fromthe initial center in Siberia during the interglacial periods. Under the influence ofthe cooling climate, they have retreated in two directions: to the refugia of Asiaand South Europe. From there they have dispersed again both to the taiga andin the Mediterranean Subregion. The Siberian–Mediterranean species have a wideecological plasticity; the category includes such species as Tettigonia viridissima,Decticus verrucivorus, and Oedipoda caerulescens, widespread in Palearctic as wellas in Bulgaria.
The only Siberian–Nearctic species by origin is Tetrix subulata with Holarcticdistribution.
The only species originating from the Mongolian center of dispersal isEpacromius tergestinus. The Asian part of its range resembles that of the Siberianspecies but is located more southward and does not include Siberia. The localitiesin the Balkan Peninsula are relict ones: Istria, Dobrudja (Dobrogea) between theDanube Delta and Varna, and Petrich in the Struma Valley. Elsewhere in Europe, thespecies occurs also in relict localities between Santander and Venice and betweenAugsburg and Rome.
Another species of the same genus is also found in Europe in the relict localities:Epacromius coerulipes, a Siberian faunal element with Eurosiberian distribution. Itoccurs also locally between Venice and Hungary, and within the Balkan Peninsulait is found only in the Black Sea coast of Romania and Bulgaria (Sozopol). Thehabitats inhabited by this species in Southeast Europe indicate that E. coerulipesrepresents a transitional form between the species with Siberian and Caspian (ortaiga and steppe) origins. An argument for this is the fact that in Siberia this speciesinhabits both forests and steppes.
The Central European faunal elements (Fig. 2) are the old species, which arebelieved to have originated in the extra-Mediterranean part of Europe, survived theglaciations in the refugia of Central and Eastern Europe, and afterward dispersedalmost over the entire Central Europe. Barbitistes constrictus appears to be a relict,which after its dispersal southward has survived in the Balkan Peninsula only inRila, the refugium for its only population outside Central Europe. Other CentralEuropean species also practically do not enter South Europe. Barbitistes serricauda(Fig. 2) and Stenobothrus crassipes (Fig. 2) have dispersed only to the northernmostparts of the Balkan Peninsula and Ectobius sylvestris (Blattodea), Poecilimon fussi,
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Pholidoptera littoralis, and Stenobothrus stigmaticus faberi are found furthersouthward in the entire northern half of the Balkan Peninsula. More widelydistributed in South Europe are Leptophyes punctatissima and Meconemathalassinum; only in the mountains are found Polysarcus denticauda, Pholidopteragriseoaptera, and Stenobothrus rubicundulus. Some Central European elementswith restricted distribution actually represent Carpathian–Balkan species, e.g. Pholi-doptera frivaldskyi and Odontopodisma rubripes.
The scheme of dispersal for Central European–Mediterranean species is similarto that of the Siberian–Mediterranean ones. The distinction lies in their initial centerof dispersal which here is the Central European. It is assumed that, during theglaciations, the species of this category have moved to the south to occupy therefugia in the southern part of Central Europe and in the Mediterranean. Aftereach expansion part of their populations has adapted to the conditions in theMediterranean Subregion. Platycleis albopunctata grisea and Gryllus campestrisare widespread in Bulgaria except the high mountain parts, and Apterygida mediais a rare species but its distribution in Bulgaria is insufficiently studied.
The Caspian faunal elements include mainly steppe species which inhabitTranscaucasia and areas to the north of the Black Sea. The expansive species ofthis category dispersed to a varying degree into the steppe extension in the north-eastern part of the Balkan Peninsula, the eastern part of Central Europe (Hungary),and Southwestern Siberia and Kazakhstan. Onconotus servillei occurs in Bulgaria(Fig. 3) only in the northeastern plain part of the country (see below in Distribution
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and origin of selected genera). It is one of the most typical steppe species ofanimals in Bulgaria together with such mammals as Sicista subtilis (Pallas, 1773)(Rodentia: Zapodidae), Cricetus cricetus Linnaeus, 1758 (Rodentia: Cricetidae), andMustela eversmanni Lesson, 1827 (Carnivora: Mustelidae). The similar range hasthe new for Bulgaria Platycleis medvedevi, recorded for Bulgaria from the Danubein the northernmost part of this country. More widely distributed in the lowlandsof Bulgaria are Gampsocleis glabra and Celes variabilis.
In the group with the Mediterranean origin, the Pontomediterranean (includingthe Anatolian) and Holomediterranean faunal elements are approximately equallyrepresented (Table 7). Nearly all Holomediterranean elements originate from thenorthern part of the Mediterranean Subregion and inhabit uniformly the entireterritory from the Atlantomediterranean to the Pontomediterranean second-levelcenters of dispersal. Depending on the degree of the postglacial changes in the rangesof the Holomediterranean species, they are divided chorologically into stationaryand expansive ones (de Lattin, 1967). This division is relative since gradual transi-tions exist between these two groups. As the limit for the stationary species, weaccept the northern boundary of the range of Tylopsis lilifolia. Some thermophilousspecies do not reach this limit and inhabit in Bulgaria only Struma Valley, EasternRhodopes, and Black Sea coast (Fig. 4), e.g. Iris oratoria (Mantodea), Kalotermesflavicollis (Isoptera), Haploembia solieri (Embioptera), Gryllomorpha dalmatina,and Myrmecophilus myrmecophilus. The number of expansive Holomediterraneanspecies is higher than that of the stationary ones (Table 8). The expansion of Tetrixceperoi and Oecanthus pellucens reaches farthest northward.
There are no systematic groups of animals in Bulgaria which includeAdriatomediterranean faunal elements2. Although Meconema meridionale andPlatycleis stricta are expansive Adriatomediterranean species, the former is acciden-tally introduced in Bulgaria; and the latter is very likely incorrectly identifiedalmost 100 years ago (it was never found again, and no preserved material exists).Therefore, the category of Adriatomediterranean species should not be includedinto the list.
The Pontomediterranean faunal elements are also divided into stationaryand expansive ones; the expansive elements slightly predominate (Table 8).The Pontomediterranean species are also interpreted here as stationary whentheir ranges exceed the limits of the Pontomediterranean second-level centereastward in the habitats similar to the Mediterranean ones (e.g. on the BlackSea coast and in Transcaucasia), or when they are polycentric species withAdriatomediterranean–Pontomediterranean distribution but within the bounds ofthe Mediterranean Subregion, e.g. Eupholidoptera chabrieri schmidti. A fewexpansive Pontomediterranean species reach northward as far as Southern Sweden,e.g. Leptophyes albovittata. Two expansive species with ranges resembling those
2 If species have been placed to this category, their finding in Bulgaria means that they are Pontomediter-ranean or polycentric Adriatomediterranean–Pontomediterranean but not strictly Adriatomediterraneanelements.
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R o m a n i a
Iris oratoria
Dociostaurus genei
Stenonemobius bicolor ponticus
M a
c e
d o
n i
a S
e r
b i
a
T u r k e y
B l a c k S e a
Platycleis escalerai Platycleis nigrosignata Eupholidoptera chabrieri schmidti
Duroniella laticornis
Kalotermes flavicollis Haploembia solieri Arachnocephalus vestitus
28
27 28
Fig. 4 Distribution in Bulgaria of some stationary taxa of Orthopterida with Mediterranean origin.Holomediterranean faunal elements: Iris oratoria, Kalotermes flavicollis, Haploembia solieri, Arach-nocephalus vestitus, Dociostaurus genei. Pontomediterranean faunal elements: Platycleis escalerai,P. nigrosignata, Eupholidoptera chabrieri schmidti, Stenonemobius bicolor ponticus, Duroniellalaticornis.
of the Holomediterranean elements are of Pontomediterranean origin. Pterolepisgermanica undoubtedly originated from the Pontomediterranean center wherelies the main part of its range (the entire Balkan Peninsula, Anatolia, andCaucasus). It has occupied during its dispersal also the Adriatomediterranean(Italy) and Tyrrhenian (Corsica) second-level centers and has reached westwardto Southern France but does not occur in the Atlantomediterranean second-levelcenter (Iberian Peninsula). Polyphaga aegyptiaca (Blattodea) in its expansion hascrossed non-existent now land bridges connecting the Balkan Peninsula to SouthItaly, Sicily, Tunisia, and Algeria (at the same time also dispersing eastward fromthe Pontomediterranean center).
A part of the Pontomediterranean stationary species (13 species) belong to theBalkan–Anatolian chorotype (Table 6). Eight of them are placed in the category ofAnatolian origin.
Levels of centers of speciation. Analyzing the species which have evolved in theBalkan Peninsula and in the adjacent areas, i.e. the groups of the Mediterranean
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278 A. POPOV
Tab
le8
Div
isio
nof
the
Bul
gari
anor
thop
teri
dta
xaw
ithM
edite
rran
ean
and
Bal
kan
orig
inby
cent
ers
ofsp
ecia
tion.
Firs
tnu
mbe
r:th
enu
mbe
rof
taxa
orig
inat
edin
the
cent
erlis
ted
inth
efi
rst
colu
mn.
Num
ber
inpa
rent
hese
s:th
enu
mbe
rof
taxa
orig
inat
edin
the
give
nce
nter
and
inpa
rts
ofit,
i.e.
atth
esa
me
leve
lor
atlo
wer
leve
l(s)
Cen
ter
ofsp
ecia
tion
Num
ber
ofta
xa
Cen
ter
offi
rst
leve
lC
ente
rof
seco
ndle
vel
Cen
ter
ofth
ird
leve
lC
ente
rof
four
thle
vel
Cen
ter
offi
fth
leve
lC
ente
rof
sixt
hle
vel
Hol
omed
iterr
anea
n39
(159
)H
olom
edite
rran
ean
(exp
ansi
vesp
ecie
s)23
of39
Hol
omed
iterr
anea
n(s
tatio
nary
spec
ies)
16of
39Po
ntom
edite
rran
ean
30(1
20)
Pont
omed
iterr
anea
n(e
xpan
sive
spec
ies)
17of
30Po
ntom
edite
rran
ean
(sta
tiona
rysp
ecie
s)13
of30
Bal
kan–
Ana
tolia
n0
(90)
Ana
tolia
n8
Bal
kan
13(8
2)D
inar
ic8
Mon
tane
Bal
kan
9M
oesi
an1
(4)
Cen
tral
Dan
ubia
nPl
ain
and
Pred
balk
an1
Bla
ckSe
aco
ast
(nor
ther
npa
rt)
2∗
Star
aPl
anin
as.
l.1
(9)
Star
aPl
anin
aM
ts.
0(6
)W
este
rnSt
ara
Plan
ina
2C
entr
alSt
ara
Plan
ina
3E
aste
rnSt
ara
Plan
ina
1Pr
edba
lkan
see
unde
rM
oesi
anSr
edna
Gor
a1
Vito
sha
Gro
up1
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ORTHOPTERIDS OF BULGARIA 279
Rila
–Rho
dope
an1
(20)
Oso
govo
–Bel
asits
aG
roup
1(7
)O
sogo
vo1
Vla
hina
1O
graz
hden
1∗∗
Bel
asits
a3
Rila
–Pir
inG
roup
2(9
)R
ila3
Piri
n3
Slav
yank
a1
Rho
dope
s0
(3)
Wes
tern
Rho
dope
s3
Eas
tern
Rho
dope
sse
eun
der
Thr
acia
nM
aced
onia
n4
(7)
Mid
dle
Stru
ma
Val
ley
3T
hrac
ian
3(7
)E
aste
rnR
hodo
pes
1T
undj
aR
egio
n1
Stra
ndja
2B
alka
nM
edite
rran
ean
5
∗M
ost
likel
ysy
nony
ms
∗∗M
ost
likel
ya
syno
nym
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280 A. POPOV
and Balkan origin (Table 7), we can address in detail, and at the different levels,the centers of speciation for the taxa found in Bulgaria (Table 8). In many animalgroups this is not possible but the high degree of endemism in South Europe andSouthwestern Asia allows that sort of analysis for Orthoptera.
The first-level center of speciation of the species of the Mediterranean andBalkan origin is the Holomediterranean one. All 159 taxa of these have originatedin the Holomediterranean center or in its parts; for 120 of these taxa, we were ableto identify these parts more precisely. The remaining 39 taxa can be only identifiedas Holomediterranean faunal elements.
The second-level center of speciation is the Pontomediterranean center with 120taxa, 30 of which are identified as originating in this center in general, and 90, inits Balkan–Anatolian part. For some of the group of 30 species, it can be specu-lated with some certainty from which part of the Pontomediterranean center theyoriginate. For instance, Saga natoliae and Duroniella laticornis are believed to beof Anatolian origin and Pachytrachis gracilis and Paracaloptenus caloptenoides, ofBalkan origin. The Anatolian origin is assigned due to the range that occupies theentire Asian sector (Anatolia, Syria, Lebanon, Israel) and only a part of the Europeansector (the southern half of the Balkan Peninsula) of the Pontomediterranean center.Such are the ranges of Duroniella laticornis and Notostaurus anatolicus. Distri-bution of the species with a Pontomediterranean origin (the Pontomediterraneanfaunal elements) indicates that some of them are polycentric regarding their second-level center of dispersal. Their ranges cover two or more centers of second levelbut not the entire Atlantomediterranean center; therefore they are not Holomediter-ranean species. To the nine second-level centers proposed by de Lattin (1967), weadd here also the Taurian center (Crimea). de Lattin (1967) considers the southerncoast of Crimea a part of the Caspian primary center. Our interpretation of theCaspian faunal elements as steppe species, however, necessitates the separation ofSouthern Crimea as a distinct second-level center and its inclusion into the Mediter-ranean primary center. Here are some examples for polycentric Pontomediterraneanranges:
Atlantomediterranean–Tyrrhenian–Adriatomediterranean–Pontomediterranean:Pholidoptera fallax (in the Atlantomediterranean center, only in Southern France);Atlantomediterranean–Tyrrhenian – Adriatomediterranean – Pontomediterranean–Cretan: Pterolepis germanica (in the Atlantomediterranean center, only inSouthern France);Tyrrhenian–Adriatomediterranean–Pontomediterranean: Phyllodromica margi-nata (Blattodea);Adriatomediterranean–Pontomediterranean: Platycleis nigrosignata;Adriatomediterranean–Pontomediterranean–Cretan–Cyprian:Platycleis escalerai;Adriatomediterranean–Pontomediterranean–Cretan–Taurian: Empusa fasciata(Mantodea);Pontomediterranean–Cretan: Platycleis incerta;Pontomediterranean–Cyprian: Duroniella laticornis;Pontomediterranean–Taurian: Poecilimon schmidti.
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ORTHOPTERIDS OF BULGARIA 281
The third-level centers of speciation for the species distributed in Bulgaria are theBalkan and the Anatolian ones. This level includes all Bulgarian taxa, which areassigned to a certain part of the Pontomediterranean second-level center by theirorigin. The species with Balkan origin prevail in the Bulgarian orthopterid fauna(Table 8) at the third level. If the next levels (from fourth to sixth) with speciesdistributed only in the Balkan Peninsula will be added, this predominance increasesgreatly. Within the framework of the entire Pontomediterranean center, however, thenumber of the species with Anatolian origin is quite higher than the number of theBalkan species. An example for this in the Bulgarian orthopterid fauna is found ifwe examine only the species of the Balkan–Anatolian chorotype (i.e. found only inthe Balkan Peninsula and Anatolia): there are twice as many taxa with the Anatolianorigin than with the Balkan one. At third level the origin is identified also accordingto the relative sizes of the Anatolian and Balkan part of the range and according tothe origin of the related species. An example of the species with Anatolian originis Bucephaloptera bucephala (found in the entire Anatolia and the eastern part ofthe Balkan Peninsula). An example of the taxon with Balkan origin is Chorthippusporphyropterus euhedickei (found in the southern part of the Balkan Peninsulaand Northwestern Anatolia). The range of the latter subspecies is obviously notfully known since it was described 15 years ago, its correct species placement isknown only for six years, and it is hard to distinguish by the stridulatory pegs fromthe related species of biguttulus-group. Nevertheless, its origin can be accepted asBalkan because nearly all representatives of Chorthippus and most of the species ofGomphocerinae in the Bulgarian fauna have northern origin. A particular positionamong the species of Anatolian origin is occupied by Anterastes serbicus. It is amountain species with Montane Mediterranean distribution. Its origin is analyzedbelow in Distribution and origin of selected genera.
The fourth-level centers of speciation occupy specific parts of the BalkanPeninsula. There are eight such centers (Fig. 5), three of which (Dinaric, MontaneBalkan, and Balkan Mediterranean) are located outside the territory of Bulgaria.The other five centers are situated partially or entirely in Bulgaria (Table 8, Fig. 6).The Rila–Rhodopean Massif is of a paramount significance for the speciation andits number of endemic species is more than two times higher than in any of theother centers. The opposite extreme is represented by the Moesian center; it isvery likely that most or all taxa believed to have originated in it are not distinctspecies or subspecies. On the other hand, the taxa from the Balkan Mediterraneancenter are distinct. Approximately equal numbers of taxa (seven to nine) haveassigned to the other centers at the fourth level of speciation. Altogether, 82 speciesand subspecies are believed to originate from the Balkan Peninsula, includingBulgaria.
The five centers present on the territory of Bulgaria are analyzed also at the fifthand sixth levels (Table 8). The Rila–Rhodopean Massif is divided into three fifth-level centers of speciation. Most species in this massif have originated in theRila–Pirin Group and Osogovo–Belasitsa Group while the role of Rhodopes for thespeciation is considerably more restricted. The formation of the fauna of Orthoptera
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282 A. POPOV
BaMe
Mac
D i nMoBa Moe
StPl
16°
44°
20° 24° 28°
36°
40°
44°
36°
40°
16° 20° 24° 28°
ThrRi-Rh
Fig. 5 Balkan centers of origin of the orthopterid species distributed in Bulgaria. Din, Dinaric; MoBa,Montane Balkan; Moe, Moesian; StPl, Stara Planina; Ri–Rh, Rila–Rhodopean; Mac, Macedonian; Thr,Thracian; BaMe, Balkan Mediterranean. Areas above 1200 m altitude are shaded.
in Stara Planina is concentrated at the fifth level while the Predbalkan, Sredna Gora,and the mountains of the Vitosha Group have a subordinate significance.
The sixth level of speciation includes parts of certain mountain ranges or groupsin Bulgaria. Most numerous here are the taxa (three in each region), which havetheir origin in Central Stara Planina, Belasitsa, Rila, Pirin, and Western Rhodopes.The same number of species has originated in the Middle Struma Valley (fifth-levelcenter of speciation).
The determination of the speciation centers at the fourth to sixth levels is toa great extent provisory. This is not only because it is based on the geographicaldivision of Bulgaria. The probable synonymy of some taxa in the future coulddecrease the number of centers at the last two levels and the list of taxa of
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ORTHOPTERIDS OF BULGARIA 283
42
44
43
24 26 272523
E of Greenwich
44
43
42
50 km
G r e e c e
S
e r
b i a
M a
c e
d o
n i
aR o m a n i a
T u r k e y
B l a c k S e a
Pholidoptera aptera karnyi
Poecilimon orbelicus
Psorodonotus fieberi Isophya modesta longicaudata Metrioptera helleri
Poecilimon zwicki Isophya petkovi Metrioptera oblongicollis
28
24 26 272523 28
Fig. 6 Distribution in Bulgaria of some endemic taxa of Tettigonioidea with different origin. Dinaricorigin: Pholidoptera aptera karnyi. Montane Balkan origin: Psorodonotus fieberi. Moesian origin:Isophya modesta longicaudata. Stara Planina origin: Metrioptera helleri. Rila–Rhodopean origin:Poecilimon orbelicus. Macedonian origin: Poecilimon zwicki. Thracian origin: Isophya petkovi. BalkanMediterranean origin: Metrioptera oblongicollis. Areas above 1200 m altitude are shaded.
Orthoptera assigned to them. It is hard to believe that some mountain ranges suchas Vlahina and Ograzhden represent centers of speciation. No local endemics areknown from these mountains from other animal groups. For Orthoptera, this is possiblyalso true for such mountain regions as Vitosha Group, Sredna Gora, Osogovo, andEastern Stara Planina. Four subspecies of Poecilimon mistshenkoi described fromPirin, Slavyanka, Belasitsa, and from Vlahina, Maleshevska, and Ograzhden as well asthree subspecies of P. affinis described from Osogovo, Rila, and Central Sredna Gorahave to be accepted with reservations. At first sight they seem to be good examples ofongoing speciation but their status is unclear.
4·3 Distribution and origin of selected genera
The most interesting in zoogeographical respect genera of Bulgarian Orthopteridaare selected and analyzed below in detail.1. Psorodonotus (15 taxa: ten species and five additional subspecies). Range:
Balkan Peninsula (one species with three subspecies), Anatolia (four species),
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284 A. POPOV
Anatolia, Caucasus, and Transcaucasia (three species), Caucasus and Transcau-casia (two species and three additional subspecies). The genus consistsof mountain species. Center of origin of this genus is most likely theCaucasus. Psorodonotus fieberi is the only species in the Balkan Peninsula.Its three subspecies (earlier considered distinct species) are clearly distin-guished geographically: P. f. illyricus Ebner, 1923 (from Istria to Montenegro),P. f. macedonicus Ramme, 1931 (Montenegro, Kosovo, Macedonia, Albania,North and Central Greece southward to Oita Mts.), and P. f. fieberi (Serbia andBulgaria). Transitional forms between them and probably hybrids are observed.The information on the subspecies ranges needs verification. Origin of the speciesand the nominate subspecies: Montane Balkan. In Bulgaria, P. f. fieberi is foundin all high mountains (Fig. 6) from 1400 to 2200 m and in an isolated locality inEastern Stara Planina at 600 m (Pešev, 1990). Occasionally, it occurs in unusuallylarge numbers, e.g. in Central Stara Planina in 1996 (Popov et al., 2000).
2. Anterastes (ten taxa: nine species and one additional subspecies). Range:Balkan Peninsula and Anatolia (one species with two subspecies), Anatolia(nine species). The genus consists of mountain species. The origin of thegenus is undoubtedly Anatolian. Anterastes serbicus is the only speciesin the Balkan Peninsula. Range of the species: Serbia, Bulgaria, Albania,North Greece southward to Ossa Mts., Western Anatolia eastward to Aksehir(A. s. serbicus), and Macedonia (A. s. macedonicus Karaman, 1961). The status ofA. s. macedonicus needs confirmation; this subspecies most likely will be provedto be a synonym. Three species, including A. serbicus, occur in NorthwesternAnatolia (Uludag Mts.). This shows the Anatolian origin of A. serbicus. It is arelict with Montane Mediterranean (more strictly, Montane Pontomediterranean)distribution. The Montane Mediterranean species are Tertiary relicts distributedonly in the mountain areas of the Mediterranean Subregion, usually in the highestparts of the mountains (Heiss and Josifov, 1990). They are representatives ofgenera with ranges restricted only to the Mediterranean or also to the CentralAsian subregion, and are the only Mediterranean species which regularly occurand reproduce in the subalpine and alpine belts. These relicts are remainders ofthe ancient cold-tolerant Mediterranean fauna which inhabited the high mountainsteppes of South Europe, or also of Central Asia, at the end of the Tertiary.A. serbicus occurs in Bulgaria in all high mountains in the subalpine, alpine, andpartly in the forest belt from 1500 to 2700 m, and only on Slavyanka it is alsofound at 900 m (Pešev and Maran, 1963).
3. Onconotus (two species). Range: from Hungary and Serbia across Ukraine,southern part of the European Russia, and North Caucasus to Siberia (twospecies). The genus consists of steppe species. Origin of the genus: Caspian.O. servillei is the only species in the Balkan Peninsula. Its range correspondsto the range of the genus and reaches eastward to West Siberia. Origin of thespecies: Caspian. In Bulgaria, O. servillei is found (Fig. 3) only in the north-eastern part of the country (Dobrudja and the eastern half of the DanubianPlain), westward to Pleven (Buresch and Peschev, 1958). The other species,
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ORTHOPTERIDS OF BULGARIA 285
O. laxmanni (Pallas, 1771), occurs from Ukraine to Siberia. The genus isseparated into the distinct subfamily Onconotinae but recently an opinion wasexpressed that its status has to be reduced to a tribe in Tettigoniinae.
4. Saga (16 taxa: 14 species and two additional subspecies). Range: South Europe,southern parts of Central Europe, West Siberia (one species), Balkan Peninsula(three species, one of which with two subspecies), from the Balkan Peninsula toSyria (one species), Anatolia (four species), Eastern Mediterranean from EasternAnatolia to Israel, Transcaucasia, and Iran (five species, one of them with twosubspecies). The richest number of species is in Turkey (ten species and oneadditional subspecies, with one species only in European Turkey). The genusincludes the largest European orthopterid species, all of them thermophilous.The origin of the genus is Pontomediterranean. The more distant ancestors ofSaga are probably Afrotropical because, according to Kaltenbach (1967), theprimitive genera of Saginae, close to Saga, are found in Africa. Only two taxa ofthose found in the Balkan Peninsula are not endemic. The widest distribution inthe genus has S. pedo. Its origin is Caspian and it occurs in South Europe (withoutGreece) northward to Southern France, Switzerland, Austria, Czech Republic,Slovakia, Ukraine, Central Russia, and eastward to West Siberia. Other speciesof the genus have a Pontomediterranean origin in a broad sense with differentcenters of speciation of fourth level: S. natoliae (Pontomediterranean in a narrowsense; from Southern Croatia and Albania to South Bulgaria, NortheasternGreece, European Turkey, Anatolia, and Syria), S. hellenica (Balkan Mediter-ranean; Albania, Macedonia, Lyulin Mts. in Southwestern Bulgaria [Fig. 8],and Greece), S. rammei (Macedonian; Petrich and Popovitsa in South Bulgaria[Fig. 8], Macedonia, Greek Macedonia), S. campbelli campbelli (Macedonian;Kresna Gorge, Belasitsa, and Slavyanka in Southwestern Bulgaria [Fig. 8],Macedonia, Northeastern Greece), S. c. gracilis (Thracian; Dobrogea in EasternRomania, Eastern Rhodopes and Sakar in South Bulgaria [Figs. 7 and 8],Northern Aegean islands, European Turkey). Another species, S. rhodiensis Salfi,1929, occurs in Greece (Rhodes Island) as well as in Southwestern Anatoliabut this island does not belong geographically to the Balkan Peninsula. Amongother Mediterranean species, the expansive eastward species is S. ephippigeraephippigera Fischer de Waldheim, 1846 with the range: Israel, Syria, EasternAnatolia, Northwestern Iran, Armenia, and Caucasus. The five Pontomediter-ranean taxa (Fig. 8) found in Bulgaira are distributed only in South Bulgaria;the information of Ramme (1951) on S. natoliae at Varna is based on the veryold material; the species is probably extinct there. The Caspian element S. pedois found mainly in North Bulgaria (most or all published records from SouthBulgaria are very likely a result of incorrect identification).
5. Bradyporus (one species) and Callimenus (six species). These two genera areclosely related. Range of both genera: Eastern Romania (Romanian Moldovaand Dobrogea), Bulgaria, Serbia, Macedonia, and North Greece. Out of thisterritory, Bradyporus is found in the north in Hungary and in the southeastin European Turkey, and Callimenus in the northeast from Southern Romania
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286 A. POPOV
Fig. 7 Saga campbelli gracilis, an endemic species and subspecies from the eastern part of the BalkanPeninsula with Thracian origin (Photo: T. Ivanova).
and Moldova to North Caucasus and in the southeast from South Greece toIran. Both genera consist of steppe species. The origin of the genera, or, moreexactly, of their common ancestor, is probably Caspian (although Bradyporusdoes not currently inhabit this area). The genus Callimenus has a Pontian type ofdistribution and the ranges of its species cover various territories around theBlack Sea: C. multituberculatus (Fischer de Waldheim, 1833) is found from theNorth Caucasian steppes to Ukraine; C. montandoni Burr, 1898, from Ukraine toSouthern Romania; C. oniscus Burmeister, 1838, in Macedonia and the entireGreece (without Thrace); C. macrogaster, from Romanian Moldova, Bulgaria,and Serbia over the Greek Thrace to Anatolia; C. dilatatus Stål, 1875, in Anatolia;and C. latipes Stål, 1875, in Iran. Judging from their current distribution, theorigin of the species occurring in Bulgaria (Callimenus macrogaster, see Fig. 9,and Bradyporus dasypus) is probably Moesian. Both species occur in Bulgariain the lowlands but the abundance of C. macrogaster decreases strongly andin many localities it became extinct during the last 60 years. These two generaare separated into the distinct subfamily Bradyporinae within the family Brady-poridae, but, according to a recent opinion, the status of this group has to bereduced to the tribe Bradyporini (Tettigoniidae: Bradyporinae).
6. Gryllomorpha (32 taxa: 31 species and one additional subspecies). Range: theMediterranean countries and Uzbekistan. The ranges of the species cover: theentire Mediterranean (G. dalmatina); the Western Mediterranean from Spain toItaly (G. uclensis Pantel, 1890); Portugal and Spain (two species, one of whichalso in North Africa); Italy (one species); Albania and continental Greece (two
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ORTHOPTERIDS OF BULGARIA 287
42
44
43
23 24 2625
24 27262523E of Greenwich
44
43
42
50km
Saga rammeiSaga cf. hellenica Saga campbelli campbelli Saga campbelli gracilis
27 28
28
S
e r
b i a
M a
c e
d o
n i
aR o m a n i a
T u r k e y
B l a c k Se a
Fig. 8 Distribution in Bulgaria of the endemic species of Saga: Saga cf. hellenica, S. rammei, S. campbellicampbelli, S. c. gracilis. Areas above 1200 m altitude are shaded.
species); Aegean islands (one species); Crete (one species); Bulgaria, KhersonRegion and Crimea in Ukraine, and Uzbekistan (G. miramae); Anatolia (onespecies); Syria (one species), Israel (three species); Egypt, Libya, and Tunisia(three species); Algeria (five species); and Morocco and Canary Islands (ninespecies). The highest number of species is found in Spain (four species; sixwith the Canary Islands) and Greece (five species). The species lead a crypticmode of life and occur in litter, under stones, in caves and tunnels, burrows ofrodents, and houses, especially in cellars. Origin of the genus: Mediterranean.Perhaps some of the Balkan species will be proved to be synonyms. In Bulgaria,G. dalmatina and G. miramae are recorded so far in the Eastern Rhodopes(the latter species uncertainly; after a female only) and in Struma Valley. Mostlikely the former species occurs in all hot lowland areas in Bulgaria. Originof the species: G. dalmatina, Holomediterranean (stationary type of the range);G. miramae, probably Pontomediterranean (expansive type of the range).
7. Discoptila (13 taxa: 12 species and one additional subspecies). Range: theMediterranean from Morocco and Spain to Anatolia and Crimea. The ranges ofthe species cover: Morocco (one species), Spain (two species, one of them alsoin Crimea and with a doubtful record in Greece), Italy (two species), Montenegro(one species), continental Greece (two species), Aegean islands, including Crete
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288 A. POPOV
Fig. 9 Callimenus macrogaster, a peculiar steppe Balkan-Anatolian species occurring in lowlands(Photo: B. Petrov).
(two species, one of which with two subspecies), Bulgaria (one species), andAnatolia (one species). The highest number of species is found in Greece (fourspecies and one additional subspecies). The systematic position of the speciesin this genus has been discussed by Popov (1984). The species lead a crypticmode of life and occur in caves, under stones, in old uninhabited houses, cellars,ruins, in litter. They are rare and, except D. fragosoi (Bolivar, 1885), all are localendemics. Most species are known from a single locality and only from the typematerial. The absence of this genus in the remaining part of the Mediterraneanis probably due to its higher requirements for humidity compared to those ofGryllomorpha. An evidence for this is the higher number of the cave dwellersin Discoptila than in Gryllomorpha. Origin of the genus: North Mediterranean.D. buresi is an Eastern Bulgarian endemic so far known from Varna area andEastern Rhodopes. Origin of the species: Thracian.
8. Subfamily Pamphaginae. The Bulgarian representatives of this subfamily belongto the genera Paranocarodes and Paranocaracris. The species of these twogenera as well as of other related genera impress with their primitive morphologyand are ancient relicts, but also remnants of a thermophilous Tertiary fauna.The taxa with similar origin, ranges, and relationships are very few inBulgaria. The genus Paranocarodes includes 13 taxa (eight species and fiveadditional subspecies). Range: Southeastern Bulgaria, Northeastern Greece,European Turkey, Aegean islands, and Western Anatolia (three species and threeadditional subspecies), Anatolia (four species and two additional subspecies), andAzerbaijan (one species). The genus consists of preglacial thermophilous relictsinhabiting xerothermic habitats with quite scanty herbaceous vegetation. Centerof origin of the genus is Anatolia. Two species occur in the Balkan Peninsula:
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ORTHOPTERIDS OF BULGARIA 289
Paranocarodes chopardi (Eastern Rhodopes in Bulgaria and Greece; Fig. 10)and P. straubei. The latter has three subspecies: P. s. straubei (SoutheasternBulgaria, European Turkey, Northwestern Anatolia), P. s. serratus Uvarov, 1949(European Turkey and Anatolia), and P. s. insularis Ramme, 1951 (the islandsof Lesbos and Chios). In Greece, outside the border of the Balkan Peninsula,occurs also P. fieberi (Brunner von Wattenwyl, 1882) with two subspecies:P. f. fieberi (Samos Island and Western Anatolia with a doubtful record on Crete)and P. f. mytilenensis Ramme, 1951 (Lesbos Island). The genus Paranocaracrisincludes 23 taxa (seven species and 16 additional subspecies). Range: Bulgaria,Greece, and European Turkey (one species with two subspecies), Anatolia (fourspecies and 13 additional subspecies), Caucasus and Transcaucasia (four speciesand two additional subspecies). This genus also includes preglacial xerother-mophilous relicts inhabiting rocky and stony habitats, relatively poor in herba-ceous vegetation. Center of origin of this genus is Anatolia. Paranocaracrisbulgaricus is the only species in the Balkan Peninsula. Its two subspeciesare clearly distinguished geographically: P. b. bulgaricus (South Bulgaria, themountains of Falakron and Pangaion in Northeastern Greece, European Turkey)and P. b. flavotibialis Willemse, 1974 (the mountains of Smolikas and Olympusin Northwestern Greece). The ranges of the Bulgarian species of both genera areisolated. Origin and distribution of the species occurring in Bulgaria (Fig. 11):Paranocarodes straubei straubei, Anatolian; the southern part of the BulgarianBlack Sea coast and adjacent areas of Strandja; P. chopardi, Thracian; EasternRhodopes (Fig. 10); Paranocaracris bulgaricus bulgaricus, Rila–Rhodopean;Kresna Gorge, Central Pirin, Slavyanka, Stargach Mts., Mesta Valley, Western
Fig. 10 Paranocarodes chopardi, a local endemic species of the Eastern Rhodopes and preglacial relict,remnant of the thermophilous Tertiary fauna (Photo: S. Beshkov).
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290 A. POPOV
42
44
43
23 24 26 27 2825
24 25 27 282623E of Greenwich
44
43
42
50km
Black S
ea
G r e e c eTurkey
aibr
eS
Mac
edon
ia
R o m a n i a
Paranocarodes chopardiParanocaracris bulgaricus bulgaricus
Paranocarodes straubei straubeiParanocarodes sp.
Fig. 11 Distribution in Bulgaria of the species of Pamphaginae: Paranocarodes straubei straubei,Paranocarodes chopardi, Paranocarodes sp., Paranocaracris bulgaricus bulgaricus. Areas above1200 m altitude are shaded.
and Central Rhodopes. An isolated population at Sliven needs revision in orderto establish to which genus and species it belongs. The species occur mainlyat low altitudes but in the presence of suitable habitats they go high up in themountains: Paranocaracris b. bulgaricus up to 1850 m in Pirin and 1800 min Slavyanka, P. b. flavotibialis occurs in Smolikas Mts. (Greece) above thetimberline at 1800–2300 m, Paranocarodes straubei straubei inhabits in Bulgariaonly low altitudes but in Uludag Mts. (Anatolia) occurs up to 2000 m. The obser-vations in Bulgaria indicate that Paranocaracris bulgaricus (and perhaps otherspecies as well) is a calciphilous species which inhabits chalky substrates only.In Pirin Mts., e.g., it occurs only in the narrow strip of Proterozoic marbles,which stretches between Slavyanka Mts. and Orelyak Peak in Central Pirin, andhas not been found in the adjacent silicate areas (Popov, 1997b). SubfamilyPamphaginae includes 38 genera. Recently there have been proposals to divideit into four subfamilies: Orchaminae (four genera), Pamphaginae (25 genera),Nocarodesinae (seven genera), and Tropidaucheninae (two genera) as well as todowngrade the genus Paranocarodes in rank to a subgenus of the genus Tropi-dauchen. These proposals do not seem to be sufficiently justified. It should benoticed also that the formation of the Latin names of subfamilies Nocarodesinaeand Tropidaucheninae has not been grammatically correct.
9. Bohemanella (three taxa: one species with three subspecies). Range of themonotypic genus: northern continuous part in Scandinavia, North Russia, almostentire Siberia, North Kazakhstan, Kamchatka, Alaska, and Northwestern Canadaand southern isolated territories in Jura, the Alps, Sudetes Mts., Tatra Mts.,
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ORTHOPTERIDS OF BULGARIA 291
mountains of Kosovo, Macedonia, and Bulgaria, mountains of Southern Siberia,Altai, North Mongolia, Manchuria, and Sakhalin. The three subspecies ofthe only species Bohemanella frigida cover the following parts of the range:B. f. frigida in the northern continuous part; B. f. strandi (Fruhstorfer, 1921) inthe southern, isolated high mountain territories in Europe; and B. f. kamtschatkae(Sjöstedt, 1936) in Kamchatka. Ramme (1951) separated the genus Bohemanellafrom the taxonomically very closely related Melanoplus. Since then B. frigidasometimes has been treated as a species of Melanoplus, and sometimes as a repre-sentative of a distinct genus. The genus Melanoplus unites more than 240 species,all from North America. Therefore the origin of Melanoplus is Nearctic. Theancestors of B. frigida should have originated in Nearctic and performed alittle known type of migration from America to Asia (Popov, 1997b). The sameopinion was expressed by Vickery (1987) but later he considered incorrectly thisspecies a Palearctic immigrant in North America (Vickery, 1997). B. frigida is ayoung (glacial) relict with a typical Arctoalpine type of distribution. The northernpart of its disjunctive range covers the tundra and forest-tundra of Eurasia andnortwestern part of North America. The southern part includes the subalpineand alpine belts of mountains in Eurasia. The wide northern range indicatesthat center of origin of Bohemanella and B. frigida could be the Siberian orAlaskan tundra. The origin of B. f. strandi is southern, most likely in the Alps.In Bulgaria, the species occurs in three isolated localities: Botev Peak in CentralStara Planina, the crest between Kamenititsa and Vihren peaks in NorthernPirin, and on Slavyanka (Fig. 1), always above the timberline on an altitude of2050–2850 m. Some authors divide the subfamily Catantopinae into two subfam-ilies (Catantopinae and Melanoplinae) and the latter into tribes Melanoplini andPodismini.
Acknowledgments
I would like to thank Dr. Stoyan Beshkov, Dr. Teodora Ivanova, and MSc BoyanPetrov for the photos of Paranocarodes chopardi, Saga campbelli gracilis, andBradyporus dasypus kindly placed at my disposal and allowed for publication here.My gratitude is directed also to MSc Silvia Tosheva for her help in preparation ofthe maps.
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