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: alpopov@bulinfo.net

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.

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.

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

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

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

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

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

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

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

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

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

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

?

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

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

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

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.

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;

NB

u,N

orth

Bul

gari

an;

NC

Ba,

Nor

ther

nan

dC

entr

alB

alka

n;N

WB

a,N

orth

wes

tern

Bal

kan;

Oso

,O

sogo

vo;

Pal,

Pale

arct

ic;

PT–P

a,Pa

leot

ropi

cal–

Pale

arct

ic;

Ri–

P–S,

Rila

–Pir

in–S

lavy

anka

;SB

a,So

uthe

rnB

alka

n;SE

Eu,

Sout

heas

tern

Eur

opea

n;SE

E–W

A,

Sout

heas

tern

Eur

opea

n–W

este

rnA

sian

;SE

u,So

uth

Eur

opea

n;So

Mo,

Sofi

aM

ount

aino

us;

Stra

,St

rand

ja;

Stru

,M

iddl

eSt

rum

aV

alle

y;T

h–B

l,T

hrac

ian–

Bla

ckSe

aco

ast;

TrA

d,T

rans

adri

atic

;T

u–C

a,T

uran

ian–

Cau

casi

an;

Tu–

Me,

Tur

ania

n–M

edite

rran

ean;

Tun

,T

undj

a;V

l–M

–O,

Vla

hina

–Mal

eshe

vska

–Ogr

azhd

en;

WPa

,W

este

rnPa

lear

ctic

;W

Rh,

Wes

tern

Rho

dope

an;

WSt

,W

este

rnSt

ara

Plan

ina.

Ori

gin:

AdM

e,A

dria

tom

edite

rran

ean;

AfT

r,A

frot

ropi

cal;

Ana

,Ana

tolia

n;B

aMe,

Bal

kan

Med

iterr

anea

n;C

as,C

aspi

an;

CE

–Me,

Cen

tral

Eur

opea

n–M

edite

rran

ean;

CE

u,C

entr

alE

urop

ean;

Din

,D

inar

ic;

HoM

e/e,

Hol

omed

iterr

anea

n/ex

pans

ive;

HoM

e/s,

Hol

omed

iterr

anea

n/st

atio

nary

;Ir

a,Ir

ania

n;M

ac,

Mac

edon

ian;

MoB

a,M

onta

neB

alka

n;M

oe,

Moe

sian

;M

on,

Mon

golia

n;Pa

Tr,

Pale

otro

pica

l;Po

Me/

e,Po

ntom

edite

rran

ean/

expa

nsiv

e;Po

Me/

s,Po

ntom

edite

rran

ean/

stat

iona

ry;

Ri–

Rh,

Rila

–Rho

dope

an;

Sib,

Sibe

rian

;Si

–Me,

Sibe

rian

–Med

iterr

anea

n;Si

–Ne,

Sibe

rian

–Nea

rctic

;Si

Ti,

Sino

tibet

an;

SiT

u,Si

beri

anT

undr

al;

StPl

,St

ara

Plan

ina;

Thr

,T

hrac

ian.

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

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.

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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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.

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.

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

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,

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

258 A. POPOV

(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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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).

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:

ORTHOPTERIDS OF BULGARIA 261

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

262 A. POPOV

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

ORTHOPTERIDS OF BULGARIA 263

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

264 A. POPOV

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

ORTHOPTERIDS OF BULGARIA 265

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

266 A. POPOV

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

ORTHOPTERIDS OF BULGARIA 267

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

268 A. POPOV

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

ORTHOPTERIDS OF BULGARIA 269

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

270 A. POPOV

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,

ORTHOPTERIDS OF BULGARIA 271

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

ORTHOPTERIDS OF BULGARIA 273

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23 24 25 26 2827

24 27262523E of Greenwich

44

43

42

50 km

G r e e c eT u r k e y

R o m a n i aM

a c

e d

o n

i a

S e

r b

i aB

l a c k S e a

Phaneroptera falcata

Barbitistes serricauda

Chrysochraon dispar Euthystira brachyptera Myrmeleotettix maculatus

Pholidoptera griseoaptera Odontopodisma rubripes Stenobothrus crassipes

28

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

274 A. POPOV

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,

ORTHOPTERIDS OF BULGARIA 275

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

42

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24 27262523E of Greenwich

44

43

42

50 km

28

S e

r b

i a

M a

c e

d o

n i

a

R o m a n i a

T u r k e yG r e e c e

B l a c k S e a

Fig. 3 Distribution in Bulgaria of Onconotus servillei, a typical steppe species.

276 A. POPOV

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.

ORTHOPTERIDS OF BULGARIA 277

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24 27262523E of Greenwich

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43

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50 km

G r e e c e

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

278 A. POPOV

Tab

le8

Div

isio

nof

the

Bul

gari

anor

thop

teri

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edite

rran

ean

and

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kan

orig

inby

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ers

ofsp

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tion.

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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

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taxa

orig

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the

give

nce

nter

and

inpa

rts

ofit,

i.e.

atth

esa

me

leve

lor

atlo

wer

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l(s)

Cen

ter

ofsp

ecia

tion

Num

ber

ofta

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Cen

ter

offi

rst

leve

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ente

rof

seco

ndle

vel

Cen

ter

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ente

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four

thle

vel

Cen

ter

offi

fth

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ente

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hle

vel

Hol

omed

iterr

anea

n39

(159

)H

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ean

(exp

ansi

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ecie

s)23

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Hol

omed

iterr

anea

n(s

tatio

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ies)

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30(1

20)

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omed

iterr

anea

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spec

ies)

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ntom

edite

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ean

(sta

tiona

rysp

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s)13

of30

Bal

kan–

Ana

tolia

n0

(90)

Ana

tolia

n8

Bal

kan

13(8

2)D

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ic8

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tane

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kan

9M

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(4)

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tral

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ubia

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ain

and

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an1

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ckSe

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l.1

(9)

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aM

ts.

0(6

)W

este

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ara

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entr

alSt

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ina

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rnSt

ara

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rM

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ORTHOPTERIDS OF BULGARIA 279

Rila

–Rho

dope

an1

(20)

Oso

govo

–Bel

asits

aG

roup

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)O

sogo

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1∗∗

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–Pir

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roup

2(9

)R

ila3

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n3

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yank

a1

Rho

dope

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(3)

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tern

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dope

s3

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tern

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dope

sse

eun

der

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acia

nM

aced

onia

n4

(7)

Mid

dle

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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

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.

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

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

ORTHOPTERIDS OF BULGARIA 283

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24 26 272523

E of Greenwich

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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),

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,

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

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

ORTHOPTERIDS OF BULGARIA 287

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27 28

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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

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:

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).

290 A. POPOV

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44

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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.,

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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