First archaeogenetic results verify the mid-Holocene occurrence of
Dalmatian pelican Pelecanus crispus far out of present range
Elena A. Nikulina and Ulrich Schmölcke
E. A. Nikulina (email@example.com) and U. Schmölcke, Schleswig-Holstein State Museums Foundation Schloss Gottorf, Centre for
Baltic and Scandinavian Archaeology (ZBSA), Archaeogenetics, DE-24837 Schleswig, Germany.
e paper presents an evaluation of subfossil bird bones from archaeological and geological sites in Europe that shows that
birds of the genus Pelecanus occurred far out of their present range between 7.4 and 5.0 ka BP in the Danish archipelago.
Additionally, from other northwestern European regions mid-Holocene records of pelicans are known. However, due to
morphological similarities there are diﬃculties in species identiﬁcation. In this paper ancient DNA barcoding techniques
were used to clarify the species assignment of one of these records for the ﬁrst time. Our results show that the bone derives
from Dalmatian pelican Pelecanus crispus, a species that today breeds in 15 colonies in the eastern Mediterranean. is spe-
cies identiﬁcation is especially remarkable since the Dalmatian pelican is known to be less migratory. We demonstrate that
the appearance and disappearance of pelicans in northwestern Europe coincide with climate parameters, since all records
fall within warm periods. is applies for the larger group of Danish records during the Holocene thermal maximum
in northern Europe as well as for two more groups of records from central Europe and Britain dating to 1.9 and 0.8 ka
ago. Recent ecological research on the Dalmatian pelican shows that the species seems to proﬁt from the modern climate
changes and is starting to expand its range. Our paper documents that under special circumstances short-distance migrant
birds are also able to expand their ranges to areas far outside of the former distribution areas. Finally, the Dalmatian pelican
is presented as an indicator species reﬂecting special climate conditions.
e present study demonstrates not only the possibility to recover avian DNA from at least 7 ka old bones, but the
relevance of such genetic analysis in combination with archaeological data, particularly if bones could not be assigned
to species level by morphological features. In such cases, aDNA is shown as a valuable tool for the reconstruction of the
avifauna of the past.
It has long been recognized that the geographical ranges of
vertebrate species have expanded and contracted in Europe
continuously since the end of the last glacial period and the
onset of the Holocene 11.6 ka BP (thousand years before
present). e most important faunal change, occurring in
the following 2 ka, was caused by successive reforestation
in the course of the climate warming (Hewitt 1999).
In those millennia, many animal species expanded their
ranges from Ice Age refugia in the Mediterranean area to
the north (Stewart et al. 2010). is fundamental change of
the fauna, which includes the whole boreal and temperate
avifauna, is quite well known by analyses of archaeological
remains of birds (Ericson and Tyrberg 2004, von den
Driesch and Pöllath 2010), but also by phylogeographic
analyses of single species (Liukkonen-Anttila et al. 2002,
Kvist et al. 2004, Brito 2005, Langguth et al. 2013).
Ongoing changes in climate towards the North Atlan-
tic mid-Holocene thermal optimum (Jansen et al. 2008,
Renssen et al. 2009) and consequently in vegetation (Davis
et al. 2003, Kalis et al. 2003), also allowed the immigration
of Mediterranean taxa to the northwestern European area
between ∼10 and ∼6 ka BP. Well known examples are the
European pond turtle Emys orbicularis (Sommer et al. 2007)
and several marine ﬁsh species (Enghoﬀ et al. 2007). Exotic
birds have also been recorded in archaeological animal assem-
blages of Britain, central Europe and southern Scandinavia
(cf. Stewart 2004), amongst others pygmy cormorant
Phalacrocorax pygmeus, in Britain, today ranging from south-
eastern Europe to central Asia (Cowles 1981), and specimens
of the genus Pterodroma (seabirds of subtropical and tropi-
cal Atlantic Ocean) recorded in Scotland (Serjeantson 2005)
and even Sweden (Lepiksaar 1958). Some published archae-
ological records of exotic species seem to be questionable,
in particular taking the general methodological problems in
bird bone identiﬁcation into account (Stewart 2005). Not
surprisingly, some records of exotic bird species have been
corrected by morphological re-analysis (Allen 2009). A good
example of a complete revision of the distribution history
after the implementation of genetic analyses of archaeologi-
cal material are sturgeons Acipenser sturio and A. oxyrinchus
(Chassaing et al. 2013, Popović et al. 2014). In birds,
however, no comparable studies have been published yet.
One of the bird taxa whose former range requires
revision is Pelecanus. From the 19th century onwards, in
© 2015 e Authors. Journal of Avian Biology © 2015 Nordic Society Oikos
Subject Editor: Martin Paeckert. Editor-in-Chief: omas Alerstam. Accepted 3 February 2015
Journal of Avian Biology 46: 344–351, 2015
archaeological literature there are several spreading reports
about mid-Holocene pelican ﬁnds in northwestern Europe,
but these reports have not been gathered and discussed in
detail. More than this, the morphometrical identiﬁcation of
pelican bones to species level is problematic (Hatting 1963,
Lorch 1992). Furthermore, keeping the present distribution
of the two European pelican species in mind, the possibil-
ity is not ruled out that the subfossil remains derive from
an extinct species or sub-species. A combination of classical
archaeozoological data with ancient and modern DNA data
is a reliable method to assess the taxonomic status of subfos-
sil bird bones, even when the species is extinct (Steeves et al.
In this paper we present the ﬁrst archaeogenetic analy-
ses of an ancient Pelecanus bone. We applied genetic analy-
ses to identify the species of the pelican. It is further the
aim to present a compilation and analysis of all subfossil
records of pelicans in northwest Europe. Both results will
be linked together at the end, forming a picture of the
range development of a Mediterranean bird species.
Material and methods
In accordance with our research goals, the study had two
parts. Firstly, to get knowledge about the pelican species that
occurred during the mid-Holocene in northwestern Europe,
a key study was conducted. In the key study the pelican bones
from the archaeological coastal site of Rosenhof (Hartz and
Lübke 2006, Goldhammer 2008) in northern Germany were
in focus. At this Stone Age settlement one of the most
complete mid-Holocene pelican skeletons was excavated,
which can be considered as representative for the other
ﬁndings from diverse archeological sites in the northwestern
Europe. From this skeleton, the left ulna was 14C AMS-dated
to 6232 39 yr BP (KIA-30007), accordingly 7145 40
calendar yr BP (calibrated using CalPal; Weninger et al.
2008) and the right radius was used for genetic analyses.
e aDNA study was done in the aDNA Laboratory of the
Center of Baltic and Scandinavian Archeology in Schleswig,
Germany. No avian DNA, neither recent nor ancient was
analysed in this laboratory prior to the published study.
To obtain material for DNA extraction from the studied
bone, a hand-held electric drill with a 2.35 mm ball cutter was
used. Prior to drilling, the bone surface was cleaned (washed)
with a solution containing 1% of sodium hypochlorite and
additionally irradiated for 60 min with UV light (254 nm) to
reduce surﬁcial contamination. About 0.4 g of bone powder
was sampled from the diaphysis. e material was dissolved
in a solution containing 300 ml 0.5 M (pH 8.0) EDTA,
300 ml MagNA Pure DNA Tissue Lysis Buﬀer (Roche Diag-
nostics), and 20 ml Proteinase K (20 mg ml–1). e mixture
was incubated for 24 h at 37°C. Finally, the temperature
was increased to 55°C for 2 h after addition of a fresh por-
tion of 20 ml Proteinase. e mixture was centrifuged at
3000 rpm for 1 min and 400 ml of supernatant was used
for the automated silica-based extraction with MagnaPure
Compact System and Nucleic Acid Isolation Kit I (Roche
Diagnostics). Blank extractions were processed. Addition-
ally, to test authenticity the obtained results DNA from three
sheep bones from Medieval (Haithabu) was co-extracted.
We designed and applied several universal PCR primers
for the genus Pelecanus to amplify fragments of mitochon-
drial cytochrome c oxidase subunit 1 (COI) and 12S rRNA
genes. e primer pairs that generated reproducible results
are listed in Table 1.
e PCR reaction volume was 10 ml including 2 ml of
the DNA extract. Each reaction consisted of Pfu recombi-
nant, 0.06 mM KCl, 16 mM (NH4)2SO4, 2 mM MgSO4,
1 mM dNTPs, 20 mM Tris-HCl (pH 8.8), 0.1% Triton
X-100, 0.1 mg ml–1 BSA (GeneON), and 0.5 mM of each
Ampliﬁcations were performed using the follow-
ing cycling conditions: initial denaturation at 95°C
for 3 min, followed by 50 cycles of at 95°C for 30 s,
annealing at 45°C for 30 s, extension at 72°C for 40
s, and ﬁnal extension at 72°C for 5 min. PCR prod-
ucts were one time re-ampliﬁed in 20 ml reaction vol-
ume with similar composition and cycling parameters,
but the number of cycles was reduced to 30. Blank
PCR controls, blank extractions and co-extracted sheep
DNA were implemented in the PCR reaction. Such
obtained amplicons were gel-puriﬁed and cloned using
E. coli JM107 and CloneJET PCR Cloning Kit (Fermentas),
and Roti®-Transform (Carl Roth) to prepare calcium
chloride competent bacterial cells.
Transformed E. coli were incubated at 37°C for 18 h
on agar ampicillin plates. Eight to ten colonies per plate
were transferred onto a fresh selective plate and addi-
tionally incubated at 37°C for 8–10 h. We used these
colonies to perform PCR with the primer pair pJET1.2
Forward und pJET1.2 Reverse (Fermentas). e reaction
mix was as follows: 1U Taq DNA Polymerase, 0.8 mM
dNTPs, 2 mM MgSO4, 10 mM KCl, 8 mM (NH4)2SO4,
10 mM Tris-HCl (pH 8.8) (GeneON), and 0.25 mM of
each primer. Cycling parameters were: 95°C for 3 min
followed by 25 cycles at 94°C for 30 s, 60°C for 30 s,
72°C for 30 s.
PCR products were sequenced using the Sanger method
(Sanger et al. 1977) at the Inst. of Clinical Molecular
Biology, Kiel. An average three to four clones per amplicon
were sequenced. To test the reproducibility of results, all
analyses were repeated at least three times (PCR, cloning,
e obtained sequences were analysed using DNASTAR
and the online Basic Local Alignment Search Tool (BLAST)
(Altschul et al. 1990).
In the second part of our study, starting with the data-
base ‘Holocene History of the European Vertebrate fauna’
(Benecke 1999, von den Driesch and Pöllath 2010), all
archaeological records for central and northern European
pelicans were compiled based on a careful reconsideration of
the original publications (Table 2). e database mentioned
contains information from more than 7500 Late Pleistocene
and Holocene faunal assemblages.
Table 1. PCR primer for the genus Pelecanus used in this study to amplify fragments of mitochondrial cytochrome c oxidase subunit 1 (COI)
and 12S rRNA genes.
Primer name Nucleotide sequence Fragment length in base pairs
with/without primer region
Pel-R1-(COI) AGAATGTAGTGTTTAGGTTTCGGTC 85/39
Pel-R2-(COI) AGCAGGGTCGAAGAATGTAGTG 82/38
Pel-R3-(COI) ATACGTGGGAATGCTATGTCTGG 65/20
Pel-R2-(12S) ATGTACGTGCTCCAGAGCCAG 68/24
Table 2. Holocene pelican bones out-of-range. All bones are context dated.
No. Site Date (years BP) Pelican bones Reference
1 Skateholm I 7400–5900 1, identiﬁcation uncertain (ulna) Jonsson 1988
2 Østenkær 6500–5900 5 (3 fragments of a humerus, radius, ulna) Enghoff 2011
3 Havnø 7000–5700 1 (sternum) Hatting 1963
4 Mejlgård 6500–5900 2 (kind of elements not published) Ljungar 1996
5 Todbjerg Mose uncertain (bog ﬁnd) 3 (coracoid, humerus, ulna) Hatting 1963
6 Brabrand Sø 6500–5900 1 (humerus) Hatting 1963
7 Syvhøje 6500–5900 14 (cranium, mandibula, 2 vertebrae, pelvis, scapula,
coracoid left and right, humerus left and right,
ulna, radius left and right, tibiotarsus)
8 Vedbæk 6500–5900 3 (coracoid, humerus, ulna) Hatting 1963
9 Stensballe Sund 6500–5900 1 (kind of element not published) Ljungar 1996
10 Kongemose 8400–7400 1 (ulnare) Noe-Nygaard 1995
11 Præstelyng 6500–5900 4 (ulna, ulnare, scapula, vertebra) Noe-Nygaard 1995
12 Sandhuse 5600–5000 2 fragments of sternum Hatting 1963
13 Åmose 6100–5700 1 (kind of element not published) Ljungar 1996
14 Ølby Lyng 6500–5900 1 (radius) Møhl 1971
15 Troldebjerg 5300–4800 1 (radius) Hatting 1963
16 Rosenhof 6800–6500 25 (tarsometatarsus left and right, ulna, two
phalanges of wings, 20 phalanges of legs)
17 Hüde I 5700–5100 2 (scapula, metatarsus) Boessneck 1978
18 Weinberg/Hitzacker 900–800 1 (vertebra) Boessneck 1982
19 Berlin-Köpenick 900–800 1 (coracoid) Müller 1977
20 Derenburg 5200–4800 1 (radius) Müller 1977
21 Trier 2000–1800 Schindler 1974
22 Assendelft F 2000–1800 11 (fragments of a skull and a vertebra) van Wijngaarden-Bakker 1988
23 Vlaardingen 5400–4500 7 (among others humerus, ulna; details unpublished) Clason and Prummel 1978
24 Hull uncertain (bog ﬁnd) 1 (femur) Forbes et al. 1958
25 King’s Lynn 3000–2000 3 fused vertebrae Forbes et al. 1958
26 Feltwell Fen uncertain (bog ﬁnd) 1 (humerus) Forbes et al. 1958
27 Burnt Fen uncertain (bog ﬁnd) 4 (humerus, radius, ulna, tarsometatarsus) Forbes et al. 1958
28 Cambridgeshire uncertain (bog ﬁnd) 1 (humerus) Forbes et al. 1958
29 Glastonbury 2100–1900 21 (all parts of the skeleton) of minimum ﬁve
birds Andrews 1899
In the pelican bone from Rosenhof, all four primer systems
provided reproducible ampliﬁcations, all blanks remained
negative. In PCR reaction with sheep samples there was no
signal. Reciprocally, no ampliﬁcation was obtained in the
pelican sample with application of sheep-speciﬁc primer
systems, but in reaction with DNA from sheep. Overall
34 clones were sequenced. ere was no commixture of
sequences (with exception of some single bacterial DNA and
artiﬁcial products, e.g. primer dimers or vector fragments).
e sequences resulted from three independent PCR repeti-
tions with identical results. ree fragments of COI and one
fragment of 12S genes were obtained. BLAST search showed
in all cases 100% identity between the obtained sequence
and the relevant gene fragments of Pelecanus crispus (Fig. 1).
No nucleotide variations or errors introduced by DNA
damage were detected.
All sequences are submitted to the European Nucleotide
Archive. e accession numbers are as follows: LM993796,
LM993797 and LM993798 for the COI gene and
LM993799 for 12S rRNA gene.
e inventory of all reported subfossil pelican remains
outside of the modern distribution area resulted in a list of
more than 119 bones from 29 diﬀerent sites in Britain, the
Figure 1. An alignment of the obtained sequences from the studied Pelecanus bone (designated as Pelecanus sp. from Rosenhof ) with
mitochondrial sequences of the genus Pelecanus in the following genomic regions: (1–3) cytochrome c oxidase subunit 1 (LM993796,
LM993797 and LM993798), and (4) 12S rRNA (LM993799).
Netherlands, Germany and Denmark, supplemented by an
uncertain record from southern Sweden (Table 2). e list
shows the archipelago in the western Baltic Sea as a remark-
able core area, of which alone 16 records originate (Fig. 2).
Further, seven records are from the northern European plain
and six from Britain. e number of excavated pelican bones
at the sites ranges from one to 25, but with the exception of
British Glastonbury (minimum number of specimens 5)
the collected remains can derive from single individuals.
Exclusively adults, no juveniles are recorded, again with
Glastonbury as the only exception. At this archaeological
site, in addition to the remains of adults there are also some
bones from juvenile and immature birds. Apart from some
bog ﬁnds, the majority of Pelecanus records were recovered
during excavations from anthropogenic contexts. All these
records from archaeological excavations are more or less
exactly context dated by archaeological artifacts from the
same ﬁnd layer, added by the pelican bone from the archaeo-
logical site Rosenhof in the western Baltic Sea area, which
was directly dated to 7145 40 calendar yr BP. e bone
is a part of skeleton, used for our genetic analyses. Chrono-
logically, the ﬁnds cluster in the three diﬀerent time frames:
between ∼7.4 and ∼5.0 ka BP, ∼2.0 ka BP and ﬁnally ∼0.9
ka BP (Fig. 3). e most conspicuous presence in these records
takes place between 7.4 and 5.0 ka BP, with 18 of the 29 records,
all located in the coastal area of the western Baltic Sea.
In subfossil mammal bones it has been repeatedly demon-
strated that ancient DNA is able to provide insights into the
long-term dynamics of species and of population distribu-
tions (de Bruyn et al. 2011). Successful genetic analysis of
thin-walled bird bones is more diﬃcult and therefore
very few aDNA studies about birds are published so far.
ey mainly discuss evolutionary, taxonomical, or biogeo-
graphical aspects of diﬀerent bird species of the Paciﬁc and
Australian region (ratites: Cooper et al. 2001, Bunce et al.
2005; geese: Paxinos et al. 2002; petrels: Welch et al. 2014;
eagles: Fleischer et al. 2000, Hailer et al. 2015). However,
for birds there are no studies comparable to this one that
trace historic distribution ranges using archeological data in
combination with genetic analysis.
Our genetic result gives evidence for the presence of a
short-distance migrant eastern Mediterranean bird species,
Figure 2. Present breeding colonies (stars) of the Dalmatian pelican
(Pelecanus crispus; combined after Crivelli 1996 and Deinet et al.
2013) and the geographic position of the subfossil pelican records
mentioned in Table 1 (dots). Dark blue dots: mid-Holocene
records, light blue dots: late Holocene records, grey dots: not
Figure 3. Dating of Holocene pelican (Pelecanus sp.) remains (num-
bers as in Table 2) and reconstructed area-average mean annual
temperature anomalies for north-west Europe (dark) and south-east
Europe (light) (after Davis et al. 2003, Fig. 4).
Pelecanus crispus in mid-Holocene northwestern Europe.
Probably also other subfossil ﬁnds of pelicans in this region
far outside of the present range also belong to the
Dalmatian pelican. is would be in agreement with the
identiﬁcations by some morphologists that had already
assumed that most of the subfossil pelican remains from
Denmark (Hatting 1963), England (Andrews 1899), and
Germany (Müller 1977) could derive from this species.
e proven occurrence of P. crispus far north of the
modern range is somewhat surprising: its present range is in
the eastern Mediterranean and Black Sea area, it is known to
be less migratory, and, furthermore, the few archaeological
records of pelicans in the Balkan area are all identiﬁed as
P. onocrotalus on the basis of morphologic criteria (Boev
1996, Gál 2007). Today, these two species occur syntopically
in south-eastern Europe. In light of the results of the present
study, the morphological identiﬁcation of pelican records in
the Balkan area should be reviewed by archaeogenetic tests.
Anyway, between ∼7.4 and ∼5.0 ka BP, the number of
pelican records in the western Baltic Sea area is so numerous
that they probably do not originate from vagrants but point
to a regular expansion of the species range. In this context,
the radiocarbon dated bone from the Rosenhof site is of
special interest, because its age is approximately 300 yr older
than all the other non-pelican radiocarbon dated organic
remains and their archaeological context (Hartz and Lübke
2006, Table 1). According to Andrén et al. (2000), due to
the marine reservoir eﬀect a correction of 300 yr should be
calculated for the remains of animals with marine diet depos-
ited during the mid-Holocene in the southwestern Baltic Sea
area. is congruence provides strong evidence of an exclu-
sively marine diet of the bird, and indicates a long stay in
a coastal area. Since there are no bones of juveniles found
yet, this is the only indication of a presence of pelicans in
mid-Holocene northwestern Europe during the whole year,
inclusive the breeding period.
Climatic (see below) as well as environmental conditions
should have been suitable for pelicans, especially because
the slowly rising water level in the Baltic basin produced
large reed beds in the coastal areas (Schmölcke et al. 2006).
Since there is a geographical gap between the records in
the broader North Sea area from England to Scania and
the main distribution area in the south-eastern Mediterra-
nean, the mid-Holocene population of P. crispus in Europe
seems to have been disjunct and divided into two main
An obvious correlation between the former range of
Dalmatian pelicans far north and climate develop-
ments exists. e main observed time frame of P. crispus
occurrence in northwestern Europe corresponds with the
Holocene thermal maximum in that region, a period of rela-
tively warm climate (Renssen et al. 2009). As a reconstruc-
tion of the temperature based on pollen data shows (Davis
et al. 2003), at the maximum ∼6.0 ka BP the mean tempera-
ture of the warmest month in the area newly colonized by
pelicans reached 1.5°C compared to modern values, while
the winter temperature was slightly lower (cf. Seppä et al.
2009). Annual temperatures in northern Europe rose until
6.5 ka BP, before stabilizing at the modern level (Fig. 3).
At the same time, the eastern Mediterranean, the main
distribution area of Dalmatian pelicans in Europe, was also
aﬀected by shifts in climate. During the mid-Holocene the
annual precipitation in the eastern Mediterranean was higher
than today, probably in the range of 800–1300 mm, without
summer droughts (Rossignol-Strick 1999), even if there was
a general trend towards drier climate (Roberts et al. 2011).
Contrasting to northern European conditions, the mid-
Holocene temperatures followed a diﬀerent pattern, since
during mid-Holocene the temperatures were up to 1.5°C
lower compared to present day values (Davis et al. 2003).
Also, the second and the third group of pelican records
in northern parts of Europe appear in periods with wet-
ter and warmer summers than today (Büntgen et al. 2011,
Fig. 4). Remarkably, the pelican records from these two
younger phases do not reach the Danish area anymore. ey
are located more to the south, both at the North Sea coastal
region of Britain and the Netherlands and in central Europe
(Fig. 2). Signiﬁcantly, the ancient Roman author Pliny the
Elder states in his Naturalis Historia (book 10, chapter 66)
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written at ∼50 AD that pelicans live at the northern coast of
Gallia, i.e. in the present Belgium and the Netherlands.
In this context the English site Glastonbury is of special
interest. In this settlement, in addition to the remains of
adults there are also some bones from juvenile and immature
birds, suggesting that the pelicans bred in the neighborhood
of the humans and that they were used as food (Andrews
1899). e ﬁnd of juvenile pelicans in England ∼2 ka ago is
an important diﬀerence to the older group of records in the
western Baltic Sea area.
It is not just subfossil bird remains that can demonstrate
changes in distribution ranges caused by climate warming
(Tyberg 2010). Since the 1940s there are detailed observa-
tions about modern changes in the occurrence and distri-
bution of European bird species and argumentations that
the most important factor for such developments seems to
be the present climatic changes (Kalela 1949). During the
last decades several studies have veriﬁed eﬀects of the North
Atlantic Oscillation on birds, in particular in the Mediter-
ranean (Gordo et al. 2011). However, to explain range shifts
not only an increasing annual temperature, but also a row
of biotic and abiotic as well as anthropogenic factors have
to be considered (Ławicki 2014; for pelicans cf. Jiguet et al.
e observed temporal coherence between the former
occurrence of Dalmatian pelican outside of its present range
and the climate development corresponds with special adap-
tations of the species, in particular with its breeding success.
Ecological studies of P. crispus have recently demonstrated
that the breeding success as well as the mortality of these
birds is signiﬁcantly dependant on atmospheric condi-
tions: in contrast to other pelican species, breeding suc-
cess is especially high in wet years. Consequently, increased
precipitation results in an increase in colony size due to the
enhanced survival of juvenile pelicans (Doxa et al. 2010,
2012). Because the current climate changes cause an increas-
ing number of wet and hot summers at breeding places of
P. crispus in the eastern Mediterranean, the number of breed-
ing pairs has started signiﬁcantly to increase and even several
new colonies have been established since 2010 (Deinet et al.
2013, Crivelli pers. comm.). Over the long term the current
climate change might provoke a new range expansion to the
north. In the last decades, there were occasional observations
of P. crispus in Hungary and Poland (Jiguet et al. 2008), and
the ﬁrst certiﬁed record both for Germany and Denmark
occurred in 2006 (Wegst 2008). In this perspective, the
Dalmatian pelican is an excellent example that an under-
standing of the way bird species have responded to past
climate changes has relevance for models about their reaction
to the current climate development (Stewart et al. 2010).
Acknowledgements – e authors would like to thank the following
colleagues for valuable comments and support: Alain Jean Crivelli
(Arles), Inge B. Enghoﬀ (Copenhagen), Dirk Heinrich (Flensburg),
Martyn Kennedy (Dunedin), John Meadows (Kiel), Wietske
Prummel (Groningen), Kenneth Ritchie (Schleswig), John R.
Stewart (London), and Wim J. Wolﬀ (Groningen). Special thanks
go to the Subject Editor of the journal, to Frank E. Zachos (Vienna)
who made helpful and very constructive comments that improved
the paper. We further thank the Inst. of Clinical Molecular Biology
in Kiel for providing Sanger sequencing as supported in part by the
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