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29
O m . Verh. 25, 1991: 2 9 - 4 9
Arctic, Montane and Steppe birds as Glacial relicts in the
West Palearctic
By Tommy Tyrberg
1. Introduction
The theory th a t speciation of forest birds
has taken place in isolated forest refugia
during glacial periods is w ell established
both in th e P alearctic and elsew here (e. g.
H a f f e r 1974, H a r r is o n 1982, M o r e a u 1966).
While this concept is very likely correct in
its essentials it is not verifiable by fossil evidence even in the W est P alearctic (which
has by fa r th e rich est Pleistocene avian fos­
sil record in th e W orld), due to the v irtu al
absence of P leistocene avifaunas from the
p u tativ e refuge areas, especially Southern
Spain, the Southern B alkans and Asia Mi­
nor.
It is how ever not generally ap p reciated
th a t a sim ilar process of ränge fragm entation and creation of relict populations by
th e g lacial/interglacial clim atic cycle also
affects arctic, m ontane and steppe birds,
and th a t a t least in the W est P alearctic the
fossil record of this process is fairly good.
2. The “Mammoth steppe” avifauna
It has long been know n th a t steppe conditions have prevailed over large p a rts of E urope d urin g glacial periods. This steppe
which a t tim es extended essentially u n b ro ken from the A tlantic to the M ackenzie B a­
sin of n o rth w estern C anada w as c h aracterized by a cold, dry C o n t i n e n t a l clim ate, a V e­
getation u sually dom inated by A rtem isia
and a sp ectacu lar m am m alian m egafauna.
This biom e w hich has no exact co u n te rp a rt
today is know n by a variety of nam es:
“steppe tu n d r a ”, “arctic ste p p e ”, “perig lacial ste p p e ”, “loess ste p p e ”, “arctic grass­
la n d ” and “m am m oth step p e”
The avifau n a of th e m am m oth steppe has
received m uch less atten tio n th a n th e m am mals b u t a t least in E urope it is q u ite well
know n from several h u n d red sites.
It has often been noted th a t glacial m am ­
m alian fau n as are “dish arm o n io u s” i. e.
they contain f orms w hich today are found in
q u ite different h ab itats (an extrem e exam ple is M uskox Ovibos m oschatus occuring to gether w ith Lion Felis leo). This also
applies to the avifauna. Thus am ong the gallinaceous bird s (which w ere ap p aren tly
a b u n d a n t on the m am m oth steppe) the n orm ally dom inant P tarm igan Lagopus m utus
and W illow Grouse Lagopus lagopus are
reg u larily found together w ith P artrid g e
P erd ix perdix, Quail C oturnix coturnix and
B lack Grouse Tetrao tetrix.
O ther characteristic m em bers of the
m am m oth steppe avifauna are rap to rs
(especially Golden Eagle A quila chrysaetos,
K estrel Falco tinnunculus, and Snowy Owl
N yctea scandiaca), L arks (usually S kylark
A lauda arvensis though other species also
occur) and corvids (p articularily Raven
Corvus corax, R ed-billed Chough Pyrrho-
30
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corax pyrrhocorax and A lpine Chough
Pyrrhocorax graculus). In ad d itio n to these
largely arctic-alp in e form s a nu m b er of
“step p e” and “m e d ite rra n e a n ” species are
also regularily found, exam ples being
Long-legged B uzzard B uteo rufinus, P allid
H arri er Circus macrourus, R ed-footed F alcon Falco vespertinus, A lectoris p artrid g es,
Orn. Verh. 25, 1991
M elanocorypha and Calandrella larks and
Rosy S tarling Stu rnus roseus.
There is a clear clim atic grad ien t in the
com position of the faunas, the “m ed iterra ­
n e a n ” species being m ore common in Sou­
th ern Europe, b u t there is an extensive overlap of n o rth ern and Southern form s w ith
bird s like Snowy Owl and A lectoris sp. b e­
ing regularily found in th e sam e deposits.
3. Relict Distribution Patterns
Among th e biogeographically significant
characteristics of the m am m oth steppe avifauna is th e considerable fossil evidence
th a t the presen t discontinuous ranges of a
n u m ber of b ird species in th e W est P alearctic m ay plausibly be reg ard ed as relicts of a
continuous ränge d uring th e la st (W ürm ian)
glacial period. These b ird s can be divided
into three groups based on th e ir p resen t distribution :
1. P r e d o m in a n tly a r c tic b ir d s w ith is o lated Southern m o n t a n e p o p u l a t i o n s :
P tarm igan Lagopus m utus, W illow G rouse
Lagopus lagopus, D otterel E udrom ias m orinellus and Shore L ark Erem ophila alpestris.
2. M o n t a n e s p e c ie s w ith a n u m b er of
isolated p opulations in differen t m o u n tain
ranges and in some cases also a Coastal pop u latio n along th e A tlan tic coast: Rock/
W ater P ip it A n th u s spinoletta/petrosus, Al­
pine A ccentor Prunella collaris, C itril finch
Serinus citrinella, Snow Finch M ontifringilla nivalis, R ed-billed C hough Pyrrhoco­
ra x pyrrhocorax and A lpine Chough P yrr­
hocorax graculus.
The Ring Ouzel Turdus torquatus also belongs to this group b u t th e fossil record of
this species is too questionable to w a rra n t
analysis since the european Turdus species
are hardly osteologically distinguishable
except by size.
3.
E astern s t e p p e s p e c i e s w ith an iso­
lated population in the Ib erian peninsula
an d /o r M aghreb: Long-legged B uzzard B u ­
teo rufinus, Im perial E agle A quila heliaca,
Dem oiselle crane A n th ro p o id es virgo, P intailed Sandgrouse Pterocles alchata, and
B lack-bellied Sandgrouse Pterocles orientalis.
4. The Würmian fossil record
There is some confusion ab o u t th e term s
used to designate th e la st glacial p eriod and
the period to w hich they apply. In th e conte x t of this p ap er “W ü rm ian ” is reg ard ed as
a synonym to “W eichselian”, “D ev en sian ”
and “V ald ai” and is considered to begin
w ith Isotope Stage 5d ab o u t 115 000 BP and
end w ith the Y ounger D ryas sta d ia l
c. 10 000 BP.
The fossil records in fig. 1 —14 are based
on d ata from some 850 sites w ith W ürm ian
avifaunas. The q uality of the fossil record is
very uneven geographically, in p a rticu la r
w ith regard to Passerines. The best-stu d ied
area by fa r is Southern France, other areas
w ith fairly good fossil records include Sw itzerland, Italy, H ungary and p a rts of Yugoslavia, Poland and Romania. M any rieh Pleistocene avifaunas are also know n from
G reat B ritain, G erm any and Czechoslova-
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ale arctic
kia b u t these w ere m ostly studied long ago
and badly need restudy. In the Soviet U nion
the Pleistocene avifaunas of C aucasus, Crimea and p a rts of U kraine are reasonably
well know n w hile th ere are few records
from oth er areas. Pleistocene avifaunas are
also scarce or unknow n from Scandinavia,
most of th e N o rth E uropean P lain, P ortugal
and Southern Spain, th e Southern B alkans,
Asia Minor, N o rth A frica and th e N ear East.
31
It m ust be em phasized th a t the W ürm ian
lasted for approxim ately 100 000 years and
encom passed several colder and m ilder clim atic phases and th a t the m aps include re ­
cords spanning the w hole of this tim e in te rval. The records therefore indicate the “envelope” of a species’ W ürm ian ränge ra th e r
th a n th e ränge at any one time. Most of the
avifaunas are how ever late glacial
(c. 1 5 0 0 0 -1 0 0 0 0 BP).
5. Arctic/Alpine Species
5.1 P t a r m i g a n
The P tarm ig an Lagopus m u tu s is th e clas­
sic exam ple of an arctic species w ith isolated Southern m ontane populations. There is
at presen t seven p opulations in th e W est P a ­
learctic; in the Pyrenees, the Alps, the Scottish H ighlands, the S candinavian m ountains, the n o rth e rn U rals, Iceland and
Spitzbergen. These populations, w ith the
possible exception of the last two, are completely isolated and have presum ably been
Figure 1:
Present breed in g ränge (horizontal hatching) and W ürm ian records (black dots) of Ptarm igan. — G egen­
w ä rtiges V ork om m en (Schraffur) u n d w ü r m e isz e itlic h e Nachw eise (Punkte) des Alpenschneehuhns.
© Ornithologische Gesellschaft Bayern, download unter www.biologiezentrum.at
32
so since th e end of the W ürm ian. E ach p opulatio n is usually reg ard ed as a sep arate subspecies.
The m ap (Fig. 1) show s th a t P tarm ig an
w ere quite common th ro u g h o u t th e u plands
of cen tral and Southern E urope d u rin g the
W ürm ian. In the southernm ost p a rt of the
glacial ränge it w as ap p aren tly re stric te d to
the highest m ountain ranges (the Pyrenees,
the C an tab rian m ountains, the A ppennines
and the S ta ra P lanina). The species seems to
have avoided plains. The P tarm ig an w as ap ­
p aren tly q u ite scarce in th e p la in of A qui­
tain e in southw est F rance in co n trast ot the
Willow G rouse ( M o u r e r - C h a u v ir ö 1979). In
Russia it is only found in th e foothills of the
C arp ath ian s, in th e m ountains of Crim ea
and in th e U rals and th ere is v irtu ally no
records from th e N o rth E uropean plain. De-
Orn. Verh. 25, 1991
spite the general scarcity of fossil birds from
these areas the absence of P tarm igan is
probably real since there are several records
of Willow G rouse (Fig. 2) and Black Grouse
from b o th the R ussian and the N orth E u ro ­
pean plains. This probably m eans th a t the
E uropean P tarm ig an w ere m ore or less isolated from th e p opulations in the U rals and
fu rth e r east even at the glacial m axim um .
This accords well w ith the fact th a t the Pyrenean, Alpine, S cottish and S candinavian
P tarm igan form the greyish “m u tu s” group
of subspecies w hile the birds in the U rals,
S pitzbergen and Iceland belong to the
b row ner “ru p e s tris ” group. Iceland has
presum ably been colonized from G reenland, and S pitzbergen either from G reenland or from Siberia by w ay of F ranz Josefs’
L and (Ptarm igan from G reenland appa-
Figure 2:
P resent breed in g ränge (horizontal hatching) and W ürmian records (black dots) of W illow Grouse. G egen w ärtiges V orkom m en (Schraffur) u n d w ü rm eiszeitlich e N ac h w e ise (P u n k te ) des M oorschnee­
huhns.
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33
rently still occasionally reach Iceland ( G u d 1972)). W hether the P tarm ig an colonized th e S candinavian m ountains a t the
end of the W ürm ian from central E urope
across the N orth E uropean p lain or from
B ritain across the N orth Sea is uncertain.
The distance across the then p artly dry
N orth Sea from S cotland to N orw ay is no
longer th a n from G reenland to Iceland and
the N orth Sea w as th en presum ably at least
as icebound as the D enm ark S tra it today.
P tarm igan have recently been found in deposits of M id-W ürm ian in te rsta d ia l age
(c. 30 000 BP) in Skjonghelleren Cave in W e ­
stern N orw ay ( L a r s e n et al. 1987) and the
possibility th a t th e species m ight have survived the glacial m axim um on some unglaciated refugium on the N orw egian coast
should p erhaps not be entirely discounted.
Tetrao m lokosiew iczi and the C aucasian
Snow cock Tetraogallus caucasicus w hich
have been p resen t in the C aucasus a t least
since the M iddle Pleistocene ( B a r y s h n i k o v &
C h e r e p a n o v 1985, B u r c h a k - A b r a m o v i c h 1975).
The end of the W ürm ian ap p aren tly caused a straig h tfo rw ard w ith d raw al of the
W illow Grouse n o rth w a rd across the N orth
E uropean plain w here (in co n trast to the
Ptarm igan) there are a few late glacial re ­
cords from n o rth ern G erm any, D enm ark
and Southern Sweden. D uring this process
the p opulation on the B ritish Isles w as cut
off by the N orth Sea and a d ap ted to the m a­
ritim e conditions on the h e ath e r m oorlands
of G reat B ritain and Ireland. T h at no relict
W illow Grouse survived on the E uropean
m ain land is presum ably due to the absence
of su itable low land m oor and scrub hab itat.
5.2 W i l l o w G r o u s e
5.3 D o t t e r e l
The W illow G rouse Lagopus lagopus has
only one isolated p opulation in the W est P a ­
learctic, th e scoticus subspecies in the B ri­
tish Isles. This subspecies occurs in several
separate areas b u t these are p robably not
isolated w ith the possible exception of the
S hetland birds.
The fossil record (Fig. 2) shows th a t th e
Willow G rouse w as quite common in cen tral
Europe durin g the W ürm ian. The W illow
Grouse a p p aren tly did not ränge quite as f ar
south as th e P tarm igan, since th ere are no
records south of the Pyrenees or th e Alps.
On the o th er h an d the species is found a t se­
veral sites in the R ussian and N orth E u ro ­
pean plains. Both these differences in the
W ürm ian ränge com pared to the P tarm ig an
is presum ably due to the W illow G rouse not
being dependent on up lan d h ab itat. It is n o ­
table th a t n eith er th e W illow Grouse, th e
P tarm igan or th e B lack G rouse seem to have
reached th e C aucasus though all th ree spe­
cies occurred in Crimea. Possibly they w ere
excluded by th e C aucasian Black G rouse
The Southern E uropean populations of
th e D o tterel Eudrom ias m orinellus are only
doubtfully glacial relicts. They are all sm all
(Fig. 3) and subspecifically identical to the
m ain N orth E u rasian population, and since
dottereis spend the w in ter in th e subdesert
zone of N orth A frica and the M iddle East
th e m ontane population may have been
established during the Holocene by shortened m igration. Such a process w ould be facilitated by the fact th a t D ottereis seems to
have relatively little o r t s t r e u e .
The fossil record how ever indicates th a t
th e D o tterel was w idespread on the glacial
steppes of central E urope (Fig. 3) and it is
p erh ap s equally likely th a t at least some of
th e Southern populations have persisted
since the W ürm ian. This a t least seems to be
th e m ost likely origin of the ra th e r larger
populations in the m ountains of C entral
Asia.
The tw o late glacial records from A pulia
are p robably of w intering birds. This p a rt of
Italy w as ap p arently very dry steppe or
m u n d sson
34
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Orn. Verh. 25, 1991
F igure 3:
P resent breeding ränge (horizontal hatching) and W ürm ian records (black dots) of D otterel. - G egen­
w ärtig es V orkom m en (Schraffur) u n d w ü r m eisz eitlich e Nachweise (Punkte) des Mornells.
sem i-desert durin g a t least p a rt of th e last
glaciatio n as in d icated by the presence of
e. g. sandgrouse ( C a s s o l i et al. 1979) an d it
w ould presum ably have been su itab le w inte r h a b ita t for D ottereis. The reco rd from
B inagady n e a r B aku ( B u r c h a k - A b r a m o v i c h
1975) is of broadly la st in terg lacial d a te and
p ro bably from a m ig ratin g bird.
5.4 S h o r e L a r k
All th e W est P alearctic p o p u latio n s of the
Shore L arks E rem ophila alpestris are subspecifically d istin ct w ith ssp. fla va in the
north, balcanica in SE E urope, atlas in Morocco, albigula east of th e Caspian, penicillata in p a rts of A natolia and Ira n and bicornis in Southern A natolia and Lebanon.
Some 35 oth er subspecies occur in A sia and
N o rth and South A m erica. This stro n g ten dency to form subspecies presu m ab ly in d i-
cates a high degree of isolation betw een po­
pulations.
The Shore L ark is n o t very common as a
fossil b u t there are enough records to show
th a t it w as w idely d istrib u ted in central
E urope during the W ürm ian (Fig. 4). Considering the w ide glacial d istrib u tio n it is not
im m ediately obvious w hy th ere are no relict
populations in w estern Europe. Possibly the
m ontane h a b ita ts th ere are too w et for
Shore Larks. In the S candinavian m ountain s it is norm ally confined to the drier
m ontane h ea th and th e sam e seems to be
tru e in the A tlas M ountains (pers. obs.)
The only other m em ber of the genus, Temm in ck ’s H orned L ark Erem ophila bilopha,
occurs in desert h a b ita t from Morocco to
K uw ait. It is tem pting to speculate th a t this
form is derived from a pop u latio n isolated
either in M aghreb or the N ear E ast during
some earlier interglacial.
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Figure 4:
Present breed in g ränge (horizontal hatching) and W ürm ian records (black dots) of Shore Lark. — G e­
g e n w ä rtig es V orkom m en (Schraffur) und w ü rm e isz e itlic h e N achw eise (P u n kte ) der Ohrenlerche.
6. Montane Species
6.1 R o c k / W a t e r P i p i t
The R ock/W ater P ip it com plex has
recently been sp lit into two species, the
Coastal Rock P ip it and th e m ontane W ater
Pipit ( R i s b e r g in press).
The Rock P ip its of NW Europe are usually
subdivided into th ree subspecies: petrosus
on the B ritish Isles, klein sch m id tii on the
Faeroes an d littoralis in Scandinavia w hile
the W ater P ipits in the m ountains of South­
ern E urope all belong to the nom inate A nthus s. spinoletta.
The fossil record of A n th u s spinoletta
sensu lato is ra th e r exiguous, consisting of
half a dozen records in France, th ree in E ng­
land and one in Czechoslovakia (Fig. 5).
These how ever m ostly lie well outside the
present b reed in g ränge of the species and
pro b ab ly indicate th a t the ranges of th e
Rock and W ater Pipits w ere contiguous
d u rin g the W ürm ian.
6.2 C i t r i l F i n c h
The C itril Finch Serinus citrinella has a
very lim ited ränge in the m ountains of
southw estern E urope (Fig. 6). The po p u lations on C orsica and S ardinia belong to a se­
p a ra te subspecies, corsicana.
The few W ürm ian records (Fig. 6) neatly
lin k th e existing populations in M assif Cen­
tra l an d the Pyrenees. There is also tw o
M iddle Pleistocene records from Southern
F ran ce ( M o u r e r - C h a u v i r E 1975) b u t n eith er
these n or the younger finds give any clue to
th e origin of the uniquely circum scribed
rän g e of this species.
36
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Orn. Verh. 25, 1991
F igure 5:
P resent breeding ränge (horizontal hatching) and W ürmian records (black dots) of R ock/W ater Pipit. G eg en w ä rtiges Vorkom m en (Schraffur) un d w ü r m eisz eitlich e Nachw eise (Punkte) v on S tra n d /W a ss er­
pieper.
6.3 S n o w F i n c h
The Snow Finch M ontifringilla nivalis occurs in a n um ber of isolated m ontane p o p u ­
lations from the C an tab rian M ountains to
Tibet. The E uropean p opulations belong to
M ontifringilla nivalis nivalis w hile th e
birds in C aucasus - A rm enia belong to th e
subspecies alpicola. The Snow Finch is
quite common in W ürm ian avifaunas of
Southern France and th e A lpine countries
(Fig. 7). The French sites lin k the e x ta n t po ­
pulation s in the Pyrenees and the Alps. D u ­
ring th e late G lacial th e species ap p aren tly
ranged f a rth e r n o rth in cen tral E urope th a n
today (the n o rth ern m o st finds are from
O berfranken ( B r u n n e r 1939, 1959)). The
scarcity of finds in eastern E urope —th ere is
none betw een A ustria and C rim ea — is
puzzling since th ere is a fa ir num ber of w ellstudied avifaunas from H ungary, Yugoslavia and Rom ania w here th e species m ight be
expected to occur. The subspecific Separa­
tion betw een the european populations and
th e Snow Finches in C aucasia — A rm enia
m ay how ever indicate a discontinuous distrib u tio n in so u th eastern E urope even under glacial conditions.
6.4 C h o u g h s
N ext to the P tarm ig an and the W illow
Grouse the tw o P yrrhocorax species are
am ong the com m onest and most characteristic birds in european Pleistocene avifau­
nas. U n fortunately the nom enclature of
these tw o species is extrem ely confused in
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37
Figure 6:
P resent breed in g ränge (horizontal hatching) and W ürmian records (black dots) of Citril Finch. — Ge­
ge n w ärtiges Vork om m en (Schraffur) u n d w ü r m eisz eitlich e N a chw eise (Punkte) des Z itronengirlitz.
the older lite ra tu re and in some cases it is
virtually im possible to decide w hich species
the records refer to (for some d etails on the
nom enclatu ral problem s see J ä n o s s y 1954).
6.4.1 R ed-billed Chough
Four subspecies of R ed-billed Chough
P yrrhocorax pyrrhocorax occur in the w est
Palearctic, pyrrhocorax in G reat B ritain
and Ireland, erythrorham phus in Western
Europe, barbarus in M aghreb and docilis in
eastern E urope and th e N ear East.
D uring th e W ürm ian th e R ed-billed
Chough occurred throughout Southern E uro­
pe (Fig. 8). The n o rth ern lim it of th e d istribution w as ap p aren tly the foothills of M assif C entral, th e Alps and the B alkan m oun­
tains south of the D anube. The W ürm ian
records lin k v irtu ally all ex tan t W est P ale­
arctic populatio n s and th e R ed-billed
Chough then also occurred in several areas
w here it is no longer found e. g. the Balearics, n o rth ern Yugoslavia and th e Crimea.
H ow ever in m ost areas the R ed-billed
Chough is ra th e r less common as a fossil
th a n the A lpine Chough and this is p a rticularily tru e of the Balkans.
The three records from G reat B ritain
(Fig. 8) are w ell sep arated from the other
W ürm ian records. One record (K irkdale
Cave in Y orkshire [ H a r r is o n 1980, L y d e k k e r
1891]) m ay in fact be from the E em ian (Ipsw ichian) Interglacial and in th a t case the
presence of this typically m ontane b ird in
the B ritish Isles w ould be a very interesting
p arallel to the species’ ränge d uring the
p resen t interglacial though the species does
n o t occur in Y orkshire today.
The other tw o sites (Chudleigh and Pavilan d Cave [ B e l l 1915, 1922, H a r r i s o n 1980])
are e ither of late G lacial or (more likely) Ho-
38
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Orn. Verh. 25, 1991
F igure 7:
P resent breeding ränge (horizontal hatching) and W ürm ian records (black dots) of S n ow Finch. — Ge­
g e n w ärtig es V orkom m en (Schraffur) u n d w ü r m eiszeitlich e Nachw eise (Punkte) des Schneefinks.
locene age. The R ed-billed Chough presu m ­
ably recolonized the B ritish Isles d uring the
L ate G lacial by follow ing th e A tlan tic coast
n o rth from S pain and Southern Fran ce since
it seems unlikely th a t the species could have
survived the G lacial m axim um in th e B ri­
tish Isles. It w as definitely p resen t in W ales
by the early H olocene w hen it is recorded
from P o rt E ynon Point Cave ( H a r r is o n
1987).
C arpathians. Also in co n trast to Pyrrhoco­
rax pyrrhocorax the A lpine Chough expanded n o rthw ards to th e edge of the N orth
E uropean plain d u ring the late G lacial (the
n orth ernm ost sites are in L uxem burg, T huringia and the Ojcow region of Southern Polan d ( B o c h e n s k i 1974, F e r r a n t & F r i a n t 1940,
M u s i l 1980). The differences in th e W ürm ian
ranges of the tw o P yrrhocorax species may
be due to a greater tolerance for cold Conti­
n en tal clim ates by the A lpine Chough.
6.4.2 A lpine Chough
All european populations of the A lpine
Chough Pyrrhocorax graculus belong to the
nom inate subspecies. The A lpine Chough is
even m ore comm on th a n the R ed-billed
Chough in W ürm ian avifaunas (Fig 9). However, in co n trast to the la tte r species the
A lpine Chough was also com mon in hilly
areas in the P an n o n ian B asin and in the
6.5 A lp in e A c c e n t o r
Three subspecies of The A lpine A ccentor
Prunella collaris occur in the W est P alearctic. The nom inate subspecies occurs in SW
Europe and M aghreb, subalpina in SE Euro­
pe, Crete and W A natolia and m ontana in
Caucasus and Iran.
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39
F igure 8:
Present breed in g ränge (horizontal hatching) and W ürm ian records (black dots) of R ed -b illed Chough.
- G egen w ärtig es V orkom m en (Schraffur) un d w ü r m e isz e itlic h e Nachweise (Punkte) der Alpen kräh e.
Most W ürm ian records are from SW E uro­
pe and lin k th e p opulations in th e Pyrenees,
M assif C entral and The Alps (Fig. 10). T here
is only two records from SE E urope, on
Crete an d K arp ath o s ( W e e s ie 1984, 1987).
The find from K arp ath o s (where the species
no longer occurs) suggests th a t the A lpine
A ccentor colonized Crete from A natolia.
6.6 O t h e r s p e c i e s
In ad d itio n to th e species m entioned
above some oth er species w hich lack a fossil
record have sim ilar d istrib u tio n p attern s:
The W allcreeper Tichodrom a m uraria and
the Tw ite A ca n th is flavirostris clearly b e­
long to th e species w ith “type 2 ” d iscontinuous m o n tan e ranges, though the Tw ite is
u nusual in lacking any m ontane p o p u la ­
tions betw een eastern A natolia and the la rgely coastal subspecies in NW E urope and
also in having a low land subspecies on the
T ran scaspian steppes.
The C rim son-w inged F inch R hodopechys
sanguinea is a som ew hat anom alous case. It
occurs in the m ountains of E astern A natolia
and reap p ears in th e A tlas m ountains. As
p o in ted out by M o r e a u (1966) it is unlikely
th a t this strictly m ontane b ird could ever
have crossed the low land deserts of E gypt
and L ibya and it m ust thus be presum ed to
have colonized the A tlas around th e n o rth
side of th e M editerranean, though it has left
no trace either in the form of fossils or relict
populations. The Rock Pigeon Columba livia m ay also originally have been a “type 2”
species, b u t in this case the original ränge
h as been b lu rre d by th e presence of feral po­
pulations.
40
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Orn. Verh. 25, 1991
Figure 9:
P resent breeding ränge (horizontal hatching) and W ürm ian records (black dots) of A lpin e Chough. G egen w ärtig es Vorkom m en (Schraffur) u n d w ü r m eisz eitlich e Nachw eise (Punkte) d e r A lpendohle.
7. Steppe Birds
7.1 L o n g - l e g g e d B u z z a r d
Two subspecies of th e Long-legged B uz­
zard occur in th e W est P alearctic, B uteo rufin u s cirtensis in M aghreb and L ibya and
th e nom inate subspecies from th e B alkans
eastw ard . The Long-legged B uzzard is not
common as a W ürm ian fossil (Fig. 11).
There are single records from A zerbaijan,
H ungary and L uxem burg and a group of six
sites in southw estern France.
This d istrib u tio n of finds pro b ab ly indicates th a t the Long-legged B uzzard occured
on the m am m oth steppe across E urope, perhaps w ith a concentration in SW France.
The concentration of sites in th e p lain of
A quitain e is parallelled in an o th er steppe
species, the Saiga antelope Saiga tatarica
w hich is also found a t scattered W ürm ian
sites across E urope b u t w as ap parently
q uite common in southw estern F rance du­
ring p a rt of the late W ürm ian ( D e l p e c h 1975,
1983).
7.2 I m p e r i a l E a g le
The spanish pop u latio n of the Im perial
E agle forms a well defined subspecies
A quila heliaca adalberti w hich m ay qualify
as a separate species w hile the bird s in the
rest of the species ränge from E astern E uro­
pe to Lake B aikal belong to the nom inate
subspecies.
There are a few W ürm ian records of Im ­
p erial Eagle w ell w est of its p rese n t ränge in
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Glacial Relicts in
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41
Figure 10:
Present breeding ränge (horizontal hatching) and W ürm ian records (black dots) of A lpine A ccentor. Gegenwärtiges Vorkom m en (Schraffur) un d w ü r m eisz eitlich e Nachw eise (Punkte) der Alpenbraunelle.
the Balkans. The w esternm ost are Saccopastore near Rome ( C a s s o l i 1978) and C otencher in S w itzerlan d ( J ä n o s s y 1980) (Fig. 12).
It is in terestin g to note th a t of th e six
W ürm ian records four (Saccopastore, Cotencher, K aim an L am brecht Cave in H ungary and Ripa in Romania) are of early W ür­
m ian age (>60 000 years BP), one (G rotta
Rom anelli in A pulia) is from the late G lacial
(10 000 —12 000 BP) w hile one (Teufelslukken in A ustria) cannot be closely dated. Adm ittedly th e evidence is slight, b u t it seems
likely from these dates th a t the m ain W ür­
mian expansion of the Im perial E agles’
ränge took place early during the glacial
cycle. This fits in well w ith the considerable
differentiation of the adalberti subspecies
which argues th a t this population has been
isolated for a long time.
7.3 D e m o is e lle C r a n e
The Dem oiselle Crane A nthropoides
virgo breeds m ainly in the E u rasian steppe
zone from the U kraine to M anchuria. There
is also a sm all population in E astern A nato­
lia an d another, very isolated and now al­
m ost extinct, in M aghreb. No subspecies
have been recorded.
There are only four Würmian records of
this species: Binagady near Baku, Karmalki
on the Upper Volga, Pin Hole Cave in northern England and Gorham’s Cave in Gi­
braltar (Burchaic-A bramovich 1975, Cowles
1981, E astham 1968). Unfortunately none of
the finds are well dated, but the one from
Pin Hole Cave may be of Mid-Würmian age
(Cowles 1981) and that from Gibraltar is
probably rather older (Eastham 1968, Gam­
ble 1986). Despite the sparseness of the
fossil record a wide Würmian ränge linking
42
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Orn. Verh. 25, 1991
Figure 11:
P resent breeding ränge (horizontal hatching) and W ürm ian records (black dots) of L on g-legged B uzzard. — G egen w ärtig es Vorkom m en (Schraffur) u n d w ü r m eisz eitlich e N ac h w e ise (Punkte) des A d l e r ­
bussards.
th e isolated p o p u latio n in M aghreb w ith the
m ain ränge of th e species on th e E u rasian
steppe seems likely.
w hile the other two, G ro tta della M adonna
n e a r Torre a Nave and G ro tta Rom anelli in
A pulia, are from the late G lacial.
7.4 B l a c k - b e l l i e d S a n d g r o u s e
7.5 P i n - T a i l e d S a n d g r o u s e
The nom inate subspecies P terocles o.
orientalis occurs in the Ib e ria n Peninsula,
M aghreb, Cyprus an d A natolia a n d is rep laced by ssp. arenarius from Iran an d th e Casp ian eastw ards. There are four W ürm ian re ­
cords of this species (Fig. 13). Of these th e
th ree finds from Southern Italy are th e m ost
significant from a biogeographic view point
since they are approxim ately h alfw ay b e t­
w een th e e x ta n t p o p u latio n s in A natolia
an d Tunisia. Of these th re e finds one, Torre
a Nave in C alabria, is of early W ürm ian age
The subspeciation p a tte rn in this species
is d ifferent from th a t in th e B lack-bellied
Sandgrouse. The nom inate Pterocles a. alchata occurs in Southern F rance and the
Iberian penninsula w hile ssp. caudacutus is
found in M aghreb, Libya and south-w est
Asia.
There is only one W ürm ian record of this
species, from th e la te G lacial of G ro tta Ro­
m anelli in A pulia ( C a s s o l i 1972).
The palaeoecology of th e G ro tta Rom a­
nelli avifauna form D ryas 3, the final “cold
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43
Figure 12:
Present breed in g ränge (horizontal hatching) and W ürmian records (black dots) of Im perial Eagle. —
Gegen wärtiges V orkom m en (Schraffur) u n d w ü r m eisz eitlich e Nachw eise (Punkte) des K aise radlers.
sn a p ” of th e W ürm ian has been studied by
C a s s o l i ( C a s s o l i et al. 1 9 7 9 ) . The sandgrouse
apparently lived on th e sem i-desert litto ra l
sandplains uncovered by the low glacial
sea-level. The existance of this h a b ita t m ay
have been im p o rta n t in facilitatin g the
spread of sandgrouse in the central M edite rra n e an area, both by narrow ing w a te rgaps and by providing suitable h a b ita t since
b o th B lack-bellied and P in -tailed S a n d ­
grouse generally avoid rocky and broken
country.
8. D iscussion
8.1 D i s p e r s a l
It is n o tab le th a t m ost arctic or steppe
species w hich h a d a glacial ränge extending
over m uch of E urope have left no relict populations. Som e such species are e. g. G yrfalcon Falco rusticolus, P allid H arrier Cir­
cus m acrourus, Snow y Owl N yctea scandiaca and Snow B unting P lectrophenax n i­
valis.
On the other h and there are no fossil re ­
cords of any m ontane species w hich occured
in E urope during the W ürm ian an d w hich
has failed to survive to the present. The only
possible exception w ould be M elanocoryp h a m axim a w hich is only found on th e Tib e ta n high plateau today, b u t has been rep o rted from a site in L iguria by C a s s o l i
(1980), b u t the identification is uncertain. It
w ould seem — n ot surprisingly — th a t the
44
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Orn. Verh. 25, 1991
Figure 13:
P resent breeding ränge (horizontal hatching) and W ürmian records (black dots) of B la ck -b ellied S an d ­
grouse. — G egen w ärtig es V orkom m en (Schraffur) und w ü r m eiszeitlich e Nachweise des Sandflughuhns.
m ontane species are b e tte r ad a p te d to su rvive for long periods in isolated m ontane
“isla n d s” th a n th e arctic or steppe birds.
The only real b a rrie r to dispersal of th e
“type 2 ” m ontane b irds betw een H im alaya
and th e A tlantic Ocean seems to have been
betw een Asia M inor and B alkan. S everal
m ontane b ird s have ranges th a t extend from
C entral A sia to th e m ountains of Asia M inor
or C aucasus (Snowcocks Tetraogallus sp.,
W hite-th ro ated Robin Irania gutturalis,
G üldensted ts R ed start P hoenicurus erythrogaster, R ed-fronted Serin Serinus pusillus, R addes D unnock P runella ocularis
and G reat R osefinch Carpodacus rubicilla)
w hile every m ontane b ird species w hich occurs on b o th sides of the A egean-B lack Sea
b a rrie r has a ränge th a t extends all th e w ay
to the A tlantic.
The “type 3 ” b ird s could have colonized
M aghreb and the P yrenean P eninsula either
by w ay of the glacial steppes n o rth of the
M editerranean or along th e N o rth A frican
litto ral. The n o rth ern ro u te w ould have
been open during m ost of th e W ürm ian
w hile th e h y p erarid desert of E gypt and L ibya w ould have been m ost easily traversed
d uring periods of increased p recip itatio n
eith er during the m id W ürm ian in terstad ial
com plex or during th e early H olocene clim atic optim um .
A th ird altern ativ e is a “d iag o n al” route
from the B alkan to Tunisia by w ay of South­
ern Italy-Sicily. U nder glacial conditions
d ispersal by this route w ould have been facilitated by greatly narro w ed w ater-g ap s
an d the em ergent areas w ould probably
have been suitable h a b ita t for steppe birds.
The v irtu al absence of Pleistocene avifau­
n as from N orth A frica m akes it im possible
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45
F igure 14:
P resent breeding ranges and W ürmian records of G yrfalcon (cross-hatching/black dots) and Saker (ho­
rizontal hatching/trian gles). — G egen w ärtig es V orkom m en un d w ü r m eiszeitlich e Nachw eise von Ger­
falke (Kreuzschraffur/Punkte) und Würgfalke (Horizontalschraffur/Dreiecke).
to decide th e issue one way or the oth er b u t
a t least in the cases of the Im perial Eagle
and th e D em oiselle Crane the n o rth e rn
route w ould seem to be m ore likely.
It is possible th a t several m ore species had
“type 3 ” ranges d uring the early Holocene
b u t th a t th e d istrib u tio n p a tte rn s have been
b lu rred th ro u g h adap tio n to th e artificial
“ste p p e” h a b ita t created in E urope by ag ri­
cu ltu ral activities from th e M id-H olocene
onw ards. This in p a rtic u la r applies to the
W hite S to rk Ciconia ciconia, G reat B u stard
Otis tarda and L ittle B ustard T etrax tetrax.
These th ree species w ere fairly com m on on
the G lacial steppe and today have strongholds in the E u rasian steppe zone and in th e
Ib erian peninsula, though they also occur
m ore or less w idely on färm lan d in central
Europe.
8.2 S p e c i a t i o n
It w ould seem th a t th e approxim ately
10 000 years since the W ürm ian have in ge­
n eral been insufficient for speciation to oc­
cur though A n th u s spinoletta/petrosus is
certainly a borderline case and some other
subspecies e. g. Lagopus lagopus scoticus
and Serinus citrinella corsicana m ay also be
close to species status.
On the whole the diversity is low am ong
arctic and steppe birds w hich argues th a t
speciation due to interglacial isolation is
n o t a common event in these groups. This is
p ro b ably due to the relative shortness
(1 0 0 0 0-20000 years) and low frequency
(about every 100 000 years) of interglacials.
T here is how ever a few likely cases of such
speciation in addition to E rem ophila alpe-
46
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stris/bilopha m entioned earlier. One is
P tarm igan/W illow Grouse. The W illow
G rouse has occured in E urope since th e b eginning of th e Pleistocene ( J a n o s s y 1974)
w hile the P tarm ig an show s up only tow ards
the end of th e M iddle Pleistocene ( J a n o s s y
1976, 1980, 1984, M o u r e r - C h a u v ir £: 1975). It
seems reasonable th a t th e “n e w ” species
evolved from some isolated p opulation, b u t
there is of course no in d icatio n w here this
happened.
A nother possible case ist th e G yrfalcon
Falco rusticolus and the S ak er Falco cherrug. A p ro b ab le ancestor of these closely re ­
lated species Falco antiquus is know n from
deposits of p robably p en u ltim ate glacial
age in F ran ce and H ungary ( J a n o s s y 1977,
M o u R E R -C H A u v m fi 1975) w hile d u rin g th e
W ürm ian b o th th e G yrfalcon and th e S aker
o ccurred in Europe and ap p aren tly behaved
as p a ra p a tric species (Fig. 14). This m akes it
likely th e speciation occurred d u rin g th e
E em ian in terglacial, p robably w ith th e an cestral G yrfalcons in the A rctic and the an cestral Sakers in th e E u rasian steppe zone
and th e tw o populations isolated from each
o ther by th e taiga beit.
Among m ontane b ird s th ere is a d istin ct
center of diversity in the m ountains of Cen­
Orn. Verh. 25, 1991
tra l Asia w ith e. g. some ten species each of
Prunella, M ontifringilla and Carpodacus.
E xactly w hy there has been so m uch specia­
tio n in this area is unclear, b u t th e large and
geographically com plex m ountain system
a n d th e frequently extrem e arid ity of the in ­
term o n tan e basins have probably facilita te d isolation. The m ontane avifauna of the
W est P alearctic on the o ther h and seems to
consist largely of “im m ig ran t” species
w hich have evolved in C entral Asia. Only
th e C itril Finch seems likely to have evolved
in Europe. Interestingly speciation has a p ­
p aren tly been ra th e r m ore common in the
M iddle E a st w here tw o m ontane endem ics
w ith P alearctic affiliations occur; S yrian
S erin Serinus syriacus and A rabian Accento r P runella fagani.
An in trig u in g p a ra lle ll to th e W illow
G rouse/P tarm igan species p a ir is found
am ong th e m ontane birds in the tw o P yrrho­
corax species. In the Pyrrhocorax case it is
P yrrhocorax graculus w hich has occurred
in th e W est P alearctic from the E arly P lei­
stocene w hile Pyrrhocorax pyrrhocorax
only shows up tow ards the end of th e M iddle
Pleistocene. However in this case too, there
is no clue w here speciation took place.
Summary
A r c t i c , M o n t a n e a n d S t e p p e b i r d s a s G l a c i a l r e l i c t s in t h e W e s t P a l e a r c t i c
A num ber of birds w ith predom inantly arctic,
alpine or stepp e distributions have discontin uous
ranges in the W est P alearctic.
F ossil evid en ce sh ow s that in several cases the
m odern ränge is relict and a rem nant of a con tinuous ränge on the periglacial steppes of the latest
(W ürmian) glaciation.
A m ong arctic sp ecies this is defin itely so in the
cases of Ptarm igan and W illow G rouse (Fig. 1 —2)
and probable for Shore Lark and D otterel
(Fig. 3 - 4 ) .
A m ong m ontane birds w ith more or less iso la ­
ted p op u lation s in the m ountains of Central and
Sou thern Europe there is exten sive fossil e v i­
dence for a continuous low lan d d istrib u tion du­
rin g the W ürmian for S n ow Finch, R ed -b illed
Chough, A lpine Chough and A lpin e A ccentor
(Fig. 7 - 1 0 ) and som e data in d icatin g the sam e for
R ock/W ater P ip it and Citril F in ch (Fig. 5 —6). A
sim ilar history is lik ely for a num ber of other sp e­
cies w ith sim ilar ranges bu t w ith no fossil record,
e. g. W allcreeper, T w ite and C rim son-w inged
Finch.
There is also a num ber of stepp e b irds w ith iso ­
lated populations in SW Europe and /or M aghreb
i. e. L ong-legged Buzzard, Imperial Eagle, D em oise lle Crane, B lack -b ellied San dgrouse and P inta iled Sandgrouse. For these sp ecies there is also
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ale arctic
som e fossil data in d icatin g a continuous Wür­
m ian ränge (Fig. 1 1 -1 4 ) though the evid en ce is
w eaker than for m ost of the arctic or m ontane
birds.
A sim ilar pattern of ränge fragm entation and
isolation has presum ably occurred during previous interglacials. D esp ite this the diversity of
the affected groups is low in the W est P alearctic,
in d icatin g that such isolation has only excep tion ally led to speciation. This is probably due to the
relative shortness (10 000 —20 000 years) and low
frequency of interglacials.
47
There is how ever a few sp ecies-p airs in the
W est P alearctic w here sp eciation due to in terglacial isolation seem s likely, i. e. P tarm igan/W illow
Grouse, G yrfalcon/Saker, Shore Lark/T em m in ck ’s H orned Lark and R ed -b illed C hough/A lpin e Chough, though in m ost of these cases sp e ­
ciation m ay have occurred outsid e the W est P ale­
arctic. There is also tw o borderline cases of “in c ipient sp ec ia tio n ” am ong the taxa treated in this
paper (A u ila heliaca/adalberti and A n th u s spinoletta/p etrosus).
Zusammenfassung
A r k t i s c h e , m o n t a n e u n d S t e p p e n v o g e l a r t e n a ls E i s z e i t r e l i k t e in d e r
W e s tp a lä a r k tis
In der W estpaläarktis w eist eine A nzahl V ogel­
arten m it vorw iegend arktischer, alpiner oder
Stepp en verbreitu ng
unzusam m enhängende
A reale auf. D ie F ossilfu n d e bew eisen, daß in einer
R eihe von F ällen das gegen w ärtige Areal ein Re­
liktvorkom m en darstellt, w elch es ursprünglich
die E israndsteppen der letzten V ereisungsperiode
(W ürm -Eiszeit) um faßte. Für Arten m it arkti­
schem V erbreitungstyp trifft dies sicher bei A lp en- und M oorschneehuhn, w ah rsch einlich auch
bei O hrenlerche und M ornell zu.
Für m ontane A rten gibt es um fassendes fossiles
B elegm aterial für ein zusam m enhängendes T ief­
land sverbreitu ngsgebiet im F alle von S ch n ee­
fink, A lpenkrähe und A lpendohle sow ie für die
A lpenbraunelle. D iese Arten haben gegen w ärtig
ein disk ontinu ierliches, m ontanes Areal in den
G ebirgen von M ittel- und Südeuropa. A uch für
Strand/W asserpieper und für den Z itronengirlitz
zeich nen sich derartige V erhältnisse ab. Für Ar­
ten ohne fossile B elege, w ie M auerläufer, Berg­
hän flin g und R otflügelgim pel, dürfte d ieselb e Er­
klärung zutreffen.
Auch für eine R eihe von Stepp en vogelarten m it
isolierten P opulationen in Südw esteuropa un d/
oder im M aghreb, w ie b eisp ielsw eise für den A d­
lerbussard, den K aiseradler, den Jungfernkra­
nich, das Sandflughuhn und das Spießflughuhn,
belegen die F ossilfun de ein k on tin uierliches
w ü rm eiszeitlich es V erbreitungsgebiet, w en n ­
gleich die E videnz schw ächer als im F all der m on­
tanen V ogelarten ist.
Ä h nlich e M uster zw isch en eiszeitlich er A real­
zersp litterun gen verbunden m it Isolation der P o­
p u lation en , dürften w ährend der früheren V erei­
sungsperioden und Z w isch en eiszeiten en tstan ­
den sein. Aber trotz der U n tersch ied lich k eit der
b etroffenen Vogelgruppen ist die davon a b zu lei­
tende A rtaufspaltung und D iversitätszun ahm e
nur vergleich sw eise geringfü gig ausgefallen.
M öglicherw eise hängt das m it der kurzen D auer
der Z w isch en eiszeiten von nur 10 0 0 0 - 2 0 000
Jahren und ihrer geringen Frequenz des A u ftre­
tens zusam m en. Für einige A rtenpaare, w ie A lp en - und M oorschneehuhn, G er- und W ürgfalke,
O hren- und H ornlerche sow ie für A lpenkrähe
und A lpendohle, läßt sich die zw isc h e n e isze itli­
che A realau fspaltun g m it A rttrennung ann eh­
men, obgleich in diesen F ällen auch E in flü sse von
außerhalb der W estpaläarktis in B etracht zu zie ­
hen sind. D ie Z w illingsarten K aiseradler und
Sp anischer K aiseradler sow ie S tran d -(F els-) und
W asserpieper befinden sich gerade im Prozeß der
A rttrennung.
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