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First archaeogenetic results verify the mid-Holocene occurrence of Dalmatian pelican Pelecanus crispus far out of present range


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The 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. This species 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. This 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.
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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 ( 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
doi: 10.1111/jav.00652
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.
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
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)
Hatting 1963
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
Genetic analyses
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
distribution areas.
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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... These are, in most cases, species that are characterized by a spectacular habitus, i.e., they are particularly conspicuous due to their color, shape, or size. Such conspicuously alien species could theoretically have naturally (temporarily) migrated into the study area, as described in [47], but, much more often, they have been deliberately introduced by humans (Figure 3). ...
... Away from direct human context, the data situation is much worse. The appearance of a "new" species has, so far, been mostly interpreted as a response to changing environmental conditions [39,47,96], although their active role in the new habitat has been less widely studied. This need exists because the fundamental phenomenon, with all its effects, is of very current interest for both nature conservation [105,106] and the economy [107]. ...
Full-text available
During the Viking era and the Middle Ages, in the Baltic Sea area, the remains of alien animal species are found rarely but recurrently. These species, which were previously widely distributed in other regions, were originally considered “exotic” by the local people of the Baltic Sea region. Conversely, “exotic” was also used to describe the last local specimens of those indigenous species that had become very rare over time. Other categories of exotic animals can be defined: the first specimens of domesticated animals seen in an area, and mythical species whose existence was generally, but erroneously, assumed. In the present paper, the evidence of selected exotic species in the Baltic Sea area is analyzed with regard to both their cultural–historical and ecological significance. Many exotic specimens were used for social and hierarchical display, illustrating the individual’s sophistication and broad knowledge of the world, their wealth, and their supra-regional influence. As a result, before Christianization, these species became part of burial rites. At the same time, some of these species became or were already integral parts of the fauna of the Baltic Sea region. Some newly immigrated species were welcomed by the people, while others were considered pests. “New”, initially exotic, species formed the basis for the purely anthropogenic urban ecosystems that emerged during this period. Meanwhile, other, formerly common, species had become “exotic” because of their increasing rarity; when they became extinct, they left significant gaps in the biocoenoses wherein they were interconnected. These ecological effects, as well as some of the socio-cultural characteristics of exotic species, find parallels in modern times.
... The five primers that could cover the highest number of species were as follows: mCOIR2 (n = 767, 88.36%), K_Bird_F1 (n = 761, 87.67%), the redundant set VertebrateR1 and VR1 (n = 718, 82.72% each), AWCintR4 (n = 706, 81.34%), and BirdF1d (n = 688, 79.26%) (Table 1), with the first one designed based on the avian order Dinornithiformes, the third one for vertebrates in general, and the rest for all birds in general (Supplementary Table 1). On the other hand, the five primers that could cover the lowest number of sequences were as follows: AWCintF2 (Patel et al., 2010), L6615(tTyr)_COI (Sorenson et al., 1999), Pel-F2-COI (Nikulina & Schmölcke, 2015), Schutz03F (Schütz et al., 2020), and BC_392F (Ogawa et al., 2015) (n = 1, 0.12% each). There was a positive but not significant relation between the primer's degeneracy level and its coverage ratio (r = 0.11, p = 0.15). ...
... Only three primers were capable of binding to at least one species of all 31 bird orders here analyzed, AWCintR4, BirdF1d, and CO1R (Table 1), and these primers were all designed focused on covering the whole avian group (Supplementary Table 1). Seven primers could bind to sequences of a single avian order: COIaRt_F (Tavares & Baker, 2008), PsEmpCOIF (Kerr, 2010), L6615_(tTyr)_ COI (Sorenson et al., 1999), AWCintF2 (Patel et al., 2010), PelF2COI (Nikulina & Schmölcke, 2015), Schutz03F (Schütz et al., 2020), and BC_392F (Ogawa et al., 2015), with only the first two binding to more than one template sequence (Supplementary Table 4). ...
The process of molecular identification of species known as DNA barcoding consists on discriminating taxa based on cumulative differences of their DNA sequences and is widely used within animals, including birds. Finding the best genomic tools, such as primers, to reach most of the species within the studied groups and evaluating the coverage breadth and physicochemical properties of primers before testing in the laboratory, can save time and financial resources. Therefore, this work aimed to retrieve the available primers for the COI gene currently used on the molecular identification of birds and evaluate them for its coverage range, physicochemical properties, and performance on in silico PCR. Afterwards, we provide the best primer subsets to cover the highest number of avian sequences and the best individual primers for each bird order in terms of coverage breadth. Thirty-one bird orders had at least one COI sequence to serve as template, 152 primers available for assessing the COI gene were evaluated, and 118 could bind to at least one template sequence. No primer subset alone could cover all template sequences; however, when combining two subsets, the complete coverage of avian COI sequences analyzed was achieved. We were able to find optimal primer subsets for more specific taxonomic groups or for analysis involving the complete avian group, in order to guide researchers in the process of choosing the best primer set for each individual barcoding goal.
... There is paleontological evidence that Dalmatian Pelicans were present and breeding from at least the Mid-Holocene (Neolithic Age) in quite a few sites in Western Europe -the Danish Archipelago, the Netherlands, north Germany and the British Isles -where there is evidence for nesting until at least the Pre-Roman era (for details see Crivelli & Vizi 1981 and several references therein, Nikulina & Schmölcke 2015). Pelicans have been also present in Greece since antiquity but the available literature does not provide any documented evidence of breeding. ...
... After cooking them in the pouch in front of the stomach they vomit them up, so that when they are open they may pick out the flesh and eat it..." (Thompson 1895, Pollard 1977. Later on, in 77 A.D., the Roman writer and historian Pliny the Elder completed his multi-volume "Naturalis Historia", in which he noted that, in around 50 A.D., pelicans nested in the northern parts of Gallia (Bostock & Riley 1855, Nikulina & Schmölcke 2015, what is now Belgium, and the Netherlands, and also nested at the deltas of the Schelde, Rhine and Elbe rivers, but from the Roman period onwards any scientific documentation on pelicans in Europe is lacking. Myths prevailed again and Aesop's fable seems having survived well into Christianity, in both the Orthodox and Catholic Churches, first through "Physiologus", a Greek ecclesiastical natural history treatise (Stresemann 1975), written in Alexandria in the 2 nd c. ...
Evidence about the historical breeding distribution of Dalmatian Pelicans Pelecanus crispus and Great White Pelicans Pelecanus onocrotalus in Greece and an adjacent zone was sought in all historical scientific literature containing references to pelicans, starting from the antiquity. Pelicans were familiar to ancient Greeks, although the two species were not distinguished. Aristotle was the first to observe and describe their migration movements in the Balkans. Subsequently, nothing was mentioned for pelicans until the 16th century observations and writings of Pierre Belon du Mans. In mid-19th century the first scientific data for both species appeared and nesting of the Dalmatian Pelicans was noted for the period 1830- 1900 in seven wetlands at Peloponnese and south mainland Greece and six at northern Greece and adjacent areas along latitude 41oN. It is impossible to evaluate the actual status of pelicans in the region during 1900-1950 as none of the few contemporary ornithological publications mentioned anything about nesting of pelicans in Greece. The breeding range of Dalmatian Pelicans had probably shrunk dramatically, although it is unlikely that all the old colonies were exterminated. Until the end of 1970s, many of the wetlands where Dalmatian pelicans bred were drained or heavily altered and the number of known colonies was reduced from seven to three, with two of them in Greece, at Amvrakikos and Lake Mikri Prespa and one at Karavasta in Albania. Scientific and conservation work initiated in 1983 mainly by the “International Pelican Research and Conservation Project” and conservation organisations, led at the turn of the century to a manyfold increase of the Amvrakikos and the Lake Mikri Prespa Dalmatian pelican colonies, which subsequently gave rise to four new colonies in the period 2003-2015: Lake Kerkini, Messolonghi lagoons, Karla Reservoir and Lake Chimaditis. The overall breeding population rose from 100-120 pairs and two colonies in the late ‘60s to over 2100 pairs and six colonies in the end of the 2010s. Simultaneously, the Great White Pelican breeding population rose from less than 100 pairs and one colony to over 700 pairs and three colonies.
... While birds of aquatic habitats are undoubtedly dominant, different sets of waterfowl species were recorded at different piledwelling settlements (Fig. 8). The number of birds from forest habitats is also significantly higher at Usviaty IV site (layer B), and only at the sites of the of its range runs through South Karelia (Panteleev, Khrabry 2020 (Nikulina, Schmölcke 2015). The nearest nesting sites of Ardeola ralloides are located in the Northern Black Sea coast, the Lower Don and the Volga Delta. ...
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The paper presents the study of avifauna from the hunter-gatherer sites at the Dnieper-Dvina basin spanning time period from the 6th to 3rd millennia BC. A total of 669 bird bones were identified and attributed to 46 different bird taxa, representing resident and migrant birds. They belong to four habitat groups: waterfowl, forest, woodside and meadow-steppe. The dominance of waterfowl birds follows the common strategy of aquatic resources exploitation. Changes in the procurement strategies, use and symbolic meanings of birds can be envisaged. Reconstructed regional mean temperature fluctuations suggest a particular influence on breeding biology and migration patterns of different species.
... Учитывая наличие нескольких особей от отдельных видов, можно предположить изменение их ареала, что совпадает с периодом существенного потепления климата (Mroczkowska et al., 2021). Пеликаны также найдены на ряде археологических памятников среднего и позднего голоцена Центральной Европы (Nikulina, Schmölcke, 2015) и на стоянке Асавец-2 (Витебская обл.) (Никифоров, 2008). ...
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В статье представлено исследование орнитофауны на неолитических памятниках Днепро-Двинского междуречья – памятники Сертея I и II. Смена палеоэкологических условий и биотопов, археологичеcких культур и типов стоянок определили изменения в культурной и экономической модели древнего населения, повлияв и на набор орнитофауны в различные временные периоды. Выделяются 4 биотопические группы птиц: околоводные, опушечные, лесные и лугово-степные. Доминируют птицы из околоводной группы. Можно допустить, что птицы играли заметную роль в пищевом рационе весной и осенью. Палеоэкологические исследования позволяют реконструировать смену типов водоемов, что могло обусловить изменения в наборе видов птиц. Смена ареалов гнездования и миграций птиц может также свидетельствовать о палеоэкологических изменениях в 5–3 тыс. до н.э. Несмотря на распространенную практику изготовления орудий, украшений из костей и зубов животных на памятниках неолита Днепро-Двинского междуречья, кости птиц использовались достаточно редко. Практически вся коллекция датируется концом 4–3 тыс. до н.э. и представлена в основном пронизками и заготовками.
... Sur ces sites, la consommation des bovins semble avoir largement privilégié les adultes, les infantiles ne représentant qu'une proportion négligeable tandis que la part des juvéniles est inférieure à 30% des restes. Les habitats concernés sont, pour la plupart, des sites proto-urbains mais les occupations de Crest et de Tournon-sur-Isère consistent plutôt en des habitats groupés ouverts et de faible ampleur 760 . Il est notable que l'oppidum du Pègue 761 montre un profil d'abattage particulièrement proche des précédents, pour les bovins, bien que les animaux infantiles soient légèrement mieux représentés. ...
Durant les âges du Fer, la vallée du Rhône se trouve à la jonction entre le monde celtique du nord et du centre de l’Europe et le monde méditerranéen. Elle constitue donc un espace de circulation soumis à différentes influences culturelles. L’étude des faunes archéologiques issues des sites de la région permet donc d’approcher les spécificités de l’élevage, de la chasse et de la consommation carnée des populations protohistoriques, en un mot, de la relation entre elles et les populations animales. Par le prisme de l’archéozoologie, la vallée du Rhône apparaît comme un espace parfaitement intégré au reste de la Gaule. Ainsi, au premier âge du Fer, les pratiques alimentaires et d’élevage montrent une partition de la région entre l’influence des mondes nord-alpin et celtique méditerranéen. Au second, les pratiques impliquant des animaux et des produits d’origine animale présentent des évolutions, parmi lesquelles une augmentation plus ou moins forte de la part des porcs dans l’alimentation. Ces changements correspondent à ceux qui s’observent dans l’ensemble de la Gaule et qui sont sans doute en lien avec une réorganisation de la société. La morphologie des espèces domestiques et son évolution, de même que la pratique de la chasse, sont également plus ou moins similaires à celles du reste de la Gaule, du VIIIe au Ier siècle av. J.-C. L’influence des cultures grecque et italique, en matière alimentaire, semble être anecdotique, du point de vue des produits d’origine animale, les évolutions semblant davantage liées à l’organisation et à la démographie des populations. L’alimentation rhodanienne nous offre donc l’image d’une région pleinement intégrée aux cultures alimentaires celtiques et présente des évolutions moins liées à celles de l’environnement qu’à des problématiques sociales et économiques.
... Emys orbicularis requires high summer temperatures for the eggs to develop (Degerbøl & Krog 1951); no remains of the species are known from the Limfjord region but there are many finds from central and south-east Denmark. Pelicanus crispus is currently found in south-eastern Europe; its former occurrence in Denmark is seen as an indication of higher-than-present summer temperatures (Hatting1963; Nikulina & Schmölcke 2015). Several southern beetle species and terrestrial gastropod species that no longer live in Denmark were also part of the fauna (Johansen & Lynge 1917;Henriksen 1933;Johnsen & Krog 1948;Thomsen & Krog 1949). ...
The Kærven Syenite Complex (KSC) is one of the oldest felsic intrusions in the Tertiary East Greenland province. Here we update our previous description of the KSC and supply a greatly expanded and comprehensive geochemical dataset. New data allow us to present a more detailed petrogenetic model for the evolution of the KSC and to investigate the geochemical characteristics of igneous cumulates subjected to loss and, occasionally, replacement of residual liquid. The KSC comprises eleven mappable units that generally young westwards. Rock types range from quartz syenite to quartz alkali feldspar syenite and alkali feldspar granite. Individual intrusive units are relatively narrow and steep-sided and are collectively suggested to represent a ring dyke complex. Basement gneiss and gabbro host rocks have locally contaminated the oldest quartz syenite KSC unit, but most of the main part of the complex escaped significant influence from host rocks. A late suite of E–W to NE–SW striking peralkaline dykes of trachytic to phonolitic compositions intrude the KSC. Compositions of the KSC rocks span a considerable range in SiO2, 59–73 wt%. Concentrations of several elements vary widely for a given SiO2 (especially at SiO2 < 66 wt%), and variation diagrams do not suggest a single model for the evolution of the units of the complex. A cumulative origin is envisaged for several KSC units. Geochemical modelling suggests that KSC magmas were derived from more than one primary magma, and that the complex evolved through a four-stage process: fractional crystallisation in precursory magma chambers was followed by final emplacement of each unit, establishment of a crystal/melt mush, expulsion of part of the residual melt and, finally, crystallisation of the remaining melt. Trace element disequilibria between alkali feldspar and host rocks in two closely associated quartz alkali feldspar syenite units indicate that highly evolved residual melt was replaced by a less evolved melt phase. Modelling of potential parent melt compositions to the Kærven magmas suggests an origin not in the Iceland plume asthenosphere, but rather in a moderately enriched source, possibly in the continental lithosphere. The course of melt evolution by fractional crystallisation is indicated to have taken place in magma chambers at depth, and repeated rise of magma into the upper crustal magma chambers and crystallisation there formed the KSC. Based on our survey of published geochemical data, the inferred parental magmas seem to have few equivalents in the North Atlantic Igneous Province and may have been generated mainly from melting of enriched dry lithospheric mantle of possibly Archaean age.
... Emys orbicularis requires high summer temperatures for the eggs to develop (Degerbøl & Krog 1951); no remains of the species are known from the Limfjord region but there are many finds from central and south-east Denmark. Pelicanus crispus is currently found in south-eastern Europe; its former occurrence in Denmark is seen as an indication of higher-than-present summer temperatures (Hatting1963; Nikulina & Schmölcke 2015). Several southern beetle species and terrestrial gastropod species that no longer live in Denmark were also part of the fauna (Johansen & Lynge 1917;Henriksen 1933;Johnsen & Krog 1948;Thomsen & Krog 1949). ...
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This paper presents new marine evidence of Lateglacial and Holocene environmental changes in the western part of Limfjorden, and provides a review of the geological history/development of this part of northern Jylland, Denmark. Lateglacial clay without fossils is widespread in the region and is probably a glaciolacustrine deposit. Limfjorden began to form as a strait in the Early Holocene due to rising relative sea level and the oldest marine shells are dated to c. 9300 cal. years BP. We propose a new relative sealevel curve for the region based on new and published data, which appear to confirm that the relative sea-level change was not extremely rapid, which was suggested earlier. During the Mid-Holocene a wide connection existed from the western part of Limfjorden to the North Sea in the west and more narrow connections existed between Limfjorden and Skagerrak in the north. The marine fauna included several species that indicate warmer and more salty waters than at present. Gradually, the connections to the North Sea and Skagerrak closed due to long-shore sediment transport and deposition of aeolian sand combined with a fall in the relative sea level during the Middle- to Late Holocene. During the Viking Age, 800–1050 CE (Common Era), the western connection to the North Sea was still open, but around 1200 CE it was closed by a coastal sandy barrier and the western part of Limfjorden became brackish. The coastal barrier was flooded on several occasions but soon formed again. After 1825 CE the western connection from Limfjorden to the North Sea has been maintained artificially.
Species reintroductions are a core component of conservation but require critical assessment of past baselines. The Dalmatian Pelican Pelecanus crispus formerly occurred in Britain during the Holocene and there is interest in its possible reintroduction, but its regional extinction dynamics remain poorly understood. Pelican bones are known from subfossil and zooarchaeological contexts at 11 British Holocene sites in three regions (East Anglian Fens, Somerset Levels and Moors, Humber Valley). Dalmatian Pelican is positively identified from four of these sites, with other specimens only identified to genus level. Pelican populations were probably only present periodically in Britain during intervals of suitable climate and available habitat: there is little evidence for pelicans before the Bronze Age, and most sites are known or inferred to be Iron Age. There is a suggested Medieval specimen but no secure evidence of pelican presence after the Roman period. However, no specimens are directly dated, with age estimation reliant upon indirect dates or inference. Climatic niche modelling indicates that southern Britain was the northwestern limit of pelican climate tolerance under postglacial conditions, and pelicans may have been unable to persist regionally until the mid‐ or late Holocene. Considerable pelican breeding and foraging habitat might have existed until the Medieval period, but availability of suitable habitat fluctuated throughout the Holocene. Probabilistic analysis suggests possible mean extinction date estimates between ad 589 and ad 1587, highlighting uncertainty associated with indirect data, and hindering our ability to correlate pelican disappearance with putative extinction drivers (overexploitation, human modification of wetlands, or non‐anthropogenic environmental changes). Improving our understanding of conditions associated with past pelican occurrence is essential for potential reintroduction decision‐making, and direct dating of bones from all sites is needed to establish a secure chronological framework for correlating pelican presence and disappearance with regional landscape change. This review provides a robust framework for how to integrate past environmental baselines into conservation decision‐making and reintroduction feasibility assessments.
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The harp seal (Pagophilus groenlandicus), today a subarctic species with breeding populations in the White Sea, around the Jan Mayen Islands and Newfoundland, was a common pinniped species in the Baltic Sea during the mid-and late Holocene. It is puzzling how an ice dependent species could breed in the Baltic Sea during the Holocene Thermal Maximum (HTM), and it remains unclear for how long harp seals bred in the Baltic Sea and when the population became extirpated. We combined radiocarbon dating of harp seal bones with zooarchaeological, palaeoenvironmental and stable isotope data to reconstruct the harp seal occurrence in the Baltic Sea. Our study revealed two phases of harp seal presence and verifies that the first colonization and establishment of a local breeding population occurred within the HTM. We suggest that periods with very warm summers but cold winters allowed harp seals to breed on the ice. Human pressure, salinity fluctuations with consequent changes in prey availability and competition for food resources, mainly cod, resulted in physiological stress that ultimately led to a population decline and local extirpation during the first phase. The population reappeared after a long hiatus. Final extinction of the Baltic Sea harp seal coincided with the Medieval Warm Period. Our data provide insights for the first time on the combined effects of past climatic and environmental change and human pressure on seal populations and can contribute with new knowledge on ongoing discussions concerning the impacts of such effects on current arctic seal populations.
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Eleven bones from a petrel species not previously recorded in Britain have been recovered from archaeological sites in the Western and Northern Isles of Scotland. They are from three different settlements: the Udal [North Uist, Outer Hebrides], Kilellan Farm [Islay, Inner Hebrides], and Brettaness [Rousay, Orkney], and all are from deposits which date from broadly the first millennium A.D.. Ten are from a medium-sized gadfly petrel, Pterodroma sp. Comparisons with the North Atlantic petrels suggest that they are probably P. feae, a near threatened species which today breeds only on one small island off Madeira and another in the South Atlantic. Alternatively they may be from an extinct Pterodroma similar in size and shape to P. feae. The eleventh bone is a from a related but smaller unidentified species. In each case the bones were found with other bird bones among the food remains from the settlements. No species of gadfly petrel breeds around the British Isles today and the species are only rarely seen offshore, but the context and associations of the finds suggest that they were caught for food, in which case gadfly petrels must have bred around the coast of Scotland in the past
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detailed overview of the current state of the exploration of birds from the archaeological sites in Bulgaria (early Neolithic to the Middle Ages| is presented. 56 sites with a total of 5,306 pcs. bones and bone fragments are examined. Their subfossil bone remains are referred to at least 85 species of 15 orders of the modern ornithofauna of Bulgaria. The most numerous among them are the remains of domestic chicken (Gallus gallus domestica), representing 58.5 to 83.6% of remains. In different ages in different parts of the country as hunting birds have been used at least 30 species of the orders Anseriformres and Galliformes. Rarer hunting objects were bustards, cranes and columbids. The beginning of poultry farming in Bulgaria dates from VII century BC, when the earliest finds of poultry - domestic fowl from the ancient town of Kabile near Yambol. At the same age are the oldest remnants of the Colchic pheasant - a fact proving the insolvency of the widely spread concepts of introducing it into the Balkans from Transcaucasia in later historical times. In a half of the settlements were found remnants of diurnal and nocturnal raptors, most common among them were the Northern goshawk, Bonelli’s eagle and the Gryffon vulture. Some of them could be used as trained birds for falconry – saker falcon, Sparrowhawk, goshawk, etc. The 4 finds of 2 specimens of the Neolithic (about 6,000 BC) categorically proves the former spread of the black grouse on Bulgarian lands. The first evidence is provided by the study of their sub-fossil remains about the past spread and some other missing or endangered today bird species in Bulgarian nature - bearded vulture, White pelican, common crane, great bustard, Bonelli’s eagle, capercaillie, Griffon Vulture, etc.
Palaeontological studies show that three endemic procellariid seabird species became extinct on the remote island of St Helena in the South Atlantic Ocean. At least one of these, Pterodroma rupinarum Olson, 1975, is likely to have survived until human colonization of the island, although it is known only from subfossil bones. Several species of Pterodroma are distributed across the Atlantic, but the skull and bill of Pt. rupinarum were judged to be more similar to the Indo-Pacific Pterodroma rostrata group, which was recently split into the separate genus Pseudobulweria. We used ancient DNA techniques to sequence the mitochondrial cytochrome b gene of the extinct Pt. rupinarum, and conducted phylogenetic analyses to investigate the placement of this enigmatic taxon. In trees constructed using maximum likelihood and Bayesian inference, Pt. rupinarum did not group with Pseudobulweria, but instead fell within a strongly supported clade of Atlantic Pterodroma, including the endangered Black-capped [Pterodroma hasitata (Kuhl, 1820)] and Bermuda [Pterodroma cahow (Nichols & Mowbray, 1916)] petrels, as well as the Macaronesian petrels [Pterodroma madeira Mathews, 1934, Pterodroma feae (Salvadori, 1899) and Pterodroma deserta Mathews, 1934]. Pterodroma rupinarum shared a particularly close relationship with Pt. feae of the Cape Verde Islands, which is also the geographically closest species within the clade. Considering the osteological distinctiveness of Pt. rupinarum it was probably a separate species, or at least a highly diverged population that was isolated for a substantial period of time prior to its extinction. © 2013 The Linnean Society of London
During the excavation of archaeological site Grube-Rosenhof in 2001 and 2002, only one uniform endmesolithic layer could be identifi ed in the periphery of the former settlement. This stratum was accumulated between 4900 and 4400 calBC . To elucidate chronological differences inside the layer, position and distribution of artifacts were analyzed. Flint and ceramics exhibited different patterns in their vertical distribution, however it was not possible to detect a clear chronological order within the deposits or a pre-pottery horizon. Potentially, the distribution patterns are shaped by a landwards shift of the outcast area in the shallow water zone with rising sea level.