Late Pleistocene and Holocene whale remains (Cetacea) from ...
Transcript of Late Pleistocene and Holocene whale remains (Cetacea) from ...
University of Southern Denmark
Late Pleistocene and Holocene whale remains (Cetacea) from Denmark and adjacentcountries: Species, distribution, chronology, and trace element concentrations
Aaris-Sørensen, Kim; Rasmussen, Kaare Lund; Kinze, Carl; Petersen, Kaj Strand
Published in:Marine Mammal Science
DOI:10.1111/j.1748-7692.2009.00356.x
Publication date:2010
Document version:Final published version
Citation for pulished version (APA):Aaris-Sørensen, K., Rasmussen, K. L., Kinze, C., & Petersen, K. S. (2010). Late Pleistocene and Holocenewhale remains (Cetacea) from Denmark and adjacent countries: Species, distribution, chronology, and traceelement concentrations. Marine Mammal Science, 26(2), 253–281. https://doi.org/10.1111/j.1748-7692.2009.00356.x
Go to publication entry in University of Southern Denmark's Research Portal
Terms of useThis work is brought to you by the University of Southern Denmark.Unless otherwise specified it has been shared according to the terms for self-archiving.If no other license is stated, these terms apply:
• You may download this work for personal use only. • You may not further distribute the material or use it for any profit-making activity or commercial gain • You may freely distribute the URL identifying this open access versionIf you believe that this document breaches copyright please contact us providing details and we will investigate your claim.Please direct all enquiries to [email protected]
Download date: 17. Oct. 2021
MARINE MAMMAL SCIENCE, 26(2): 253–281 (April 2010)C© 2009 by the Society for Marine MammalogyDOI: 10.1111/j.1748-7692.2009.00356.x
Late Pleistocene and Holocene whaleremains (Cetacea) from Denmark and adjacent countries:
Species, distribution, chronology, and traceelement concentrations
KIM AARIS-SØRENSEN
Zoological Museum, National History Museum of Denmark,University of Copenhagen,
Universitetsparken 15, DK-2100 Copenhagen Ø, DenmarkE-mail: [email protected]
KAARE LUND RASMUSSEN
Institute of Physics and Chemistry,University of Southern Denmark,
Campusvej 55, DK-5230 Odense M, Denmark
CARL KINZE
CCKonsult,Rosenørns Alle 55 2 tv,
DK-1970 Frederiksberg C, Denmark
KAJ STRAND PETERSEN
Geological Survey of Denmark and Greenland (GEUS),Øster Voldgade 10,
DK-1350 Copenhagen K, Denmark
ABSTRACT
We describe and review the subfossil whale bones (mammalian order Cetacea)material from the southern Scandinavian area, that is, Skagerrak, Kattegat, theinner Danish waters and the southwestern Baltic Sea. Fifteen species were identifiedfrom the subfossil records of which all, except for the bowhead whale (Balaenamysticetus), have also been encountered in the modern times. Fifty-one specimenswere radiocarbon dated covering 12 of the subfossil species. The dates fell in threedistinct clusters with a few specimens before the last glacial maximum (LGM), alarge group between LGM and the Pleistocene/Holocene boundary (ca. 17.0–11.7cal. kyr BP), and another large group from ca. 8.0 cal. kyr BP onward. Seventeenof the radiocarbon dated specimens have been subjected to trace element analysisby Instrumental Neutron Activation Analysis. Cross plots of the concentrations ofFe and Zn, and Fe and Co show that it is possible to distinguish crayfish eatersfrom fish/squid eaters. This can be used as a novel and independent method for thedetermination to species of whale remains of otherwise uncertain speciation.
253
254 MARINE MAMMAL SCIENCE, VOL. 26, NO. 2, 2010
Key words: Late Pleistocene-Holocene, southern Scandinavia, whale remains,species, distribution, chronology, trace elements.
Subfossil whale bones (mammalian order Cetacea) have been found in relativelylarge numbers all over southern Scandinavia, and in Denmark alone 361 specimenshave been recorded from late Middle Weichselian, Late Weichselian, and Holocenedeposits. Due to glacier dynamics and dramatic changes in relative sea-level through-out these periods, cetacean bones can be found in tills, out washed sediments, and inraised marine shorelines and sediments.
Hitherto, no attempts have been made to review the entire material from thesouthern Scandinavian area, that is, Skagerrak, Kattegat, the inner Danish waters,and the southwestern Baltic Sea. Earlier, sporadic notes on the discovery of subfossilwhale remains, however, were provided by Nilsson (1847) in his survey of theScandinavian mammals, by Liljeborg (1861) in his account on the Swedish andNorwegian whale species, and again in 1874 in his work on the entire mammalianfauna of the two countries (Liljeborg 1874). During the end of the 19th centuryTauber (1878–1892) described whale remains in his Zoologica Danica.
The first publication to deal exclusively with subfossil remains of whales wasprovided by Liljeborg (1867), who described the discovery of major parts of a skeletonof a gray whale (Eschrichtius robustus) from marine sand and clay deposits at Graso,Roslagen (Sweden) and parts of a skeleton of a juvenile baleen whale designatedBalaena swedenborgii in marine sediments at Skara, Vastergotland (Sweden). Bothwhales were already mentioned by Liljeborg in 1861 in his general survey of theScandinavian whale fauna where he had named the gray whale Balaenoptera robustaand considered the Swedenborg whale as a distinct subfossil species. The latter viewwas supported by Aurivillius (1888), who described the discovery of another juvenile“Swedenborg whale,” this time in marine sediments at Tvaaker, Halland (Sweden).Further arguments for giving the Swedenborg whale specific rank were given byNybelin (1942, 1946) who concluded that the specimens resembled neither thebowhead (Balaena mysticetus) nor the Atlantic right whale (Eubalaena glacialis). Morerecently, Freden (1984) recommended postponing the final decision on the taxonomicstatus until more remains become available. Lepiksaar (1986) stressed that safespecies identification should only be based on individuals that are skeletally matureand therefore the Swedenborg whales should be regarded as juvenile specimens ofE. glacialis. The Graso specimen, on the other hand, is now generally accepted as theholotype of the gray whale and named E. robustus (Liljeborg).
The first surveys of whale remains covering a larger geographical region werecarried out by Winge (1899, 1904). These catalogs enumerate 25 Danish finds ofwhale bones belonging to bowhead whale (B. mysticetus), humpback whale (Megapteranovaeangliae), sperm whale (Physeter macrocephalus), beluga whale (Delphinapterus leu-cas), killer whale (Orcinus orca), bottlenose dolphin (Tursiops truncatus), white-beakeddolphin (Lagenorhynchus albirostris), and harbor porpoise (Phocoena phocoena) and listboth locality and other relevant details such as a tentative geological dating. Thenext overview was provided by Degerbøl (1933) in an introduction to a compre-hensive work on the prehistoric Danish mammal fauna. It added two new whalespecies, the blue whale (Balaenoptera musculus) from late glacial marine sedimentsin northern Jylland and the false killer whale (Pseudorca crassidens) from Kold-ing Fjord, South Jylland (these latter remains probably originate from the 1861
AARIS-SØRENSEN ET AL.: SUBFOSSIL WHALE BONES 255
invasion of a large school of this species into the inner Danish waters, see Kinze2007).
General faunal history reviews by Aaris-Sørensen (1998, 2007), Lepiksaar (1986),Liljegren (1975), and Liljegren and Lageras (1993) have treated finds of more recentdates and more specific contributions have been given by Lepiksaar (1966), dealingwith modern and fossil occurrences of toothed whales in Sweden, by Møhl (1970),dealing with prehistoric seal and whale hunting in Denmark, and finally by Freden(1975, 1984) the first to publish a series of radiocarbon dates of subfossil Swedishwhale remains.
The objective of this article is to present and characterize the Late Pleistoceneand Holocene whale fauna in southern Scandinavia. Successful dating of a largenumber of subfossil whale remains allows us to describe the changes in speciesrichness and composition and changes in geographical and chronological distribution.These changes can be compared with the changes in climate as reflected in thehighly dynamic environmental history of the last glacial-interglacial cycle in southernScandinavia.
It could be asked to what degree did the dramatic changes in land/sea configura-tion, water temperature, and productivity determine the composition of the marinemammal populations. Questions like this have direct relevance to modern natureconservation and biodiversity management, especially in relation to future climatechanges. The results are furthermore linked to and compared with the historicalrecords built on stranding lists and whale observations in the area for the period1575–2007 (Kinze 1995, 2007).
The background material for this study is provided mainly by the collection ofsubfossil whale bones housed at the Zoological Museum (National History Museumof Denmark), University of Copenhagen (ZMUC), as well as by the results of arecently completed dating program including radiocarbon datings of 50 whales fromthe Danish waters, the Swedish west coast, and the German Baltic coast. For a selectedgroup of specimens we have also performed trace element analysis, which may reflectthe food intake of the animal, and which in any event, helps to group less specificspecimens.
MATERIALS AND METHODS
The investigations presented here are mainly based on the collection of subfossilwhales kept at the Zoological Museum, University of Copenhagen, for the presentcomprising 361 specimens from Danish waters. Among these 41 were selected forradiocarbon dating along with seven Swedish and two German samples, which wherekindly made available to us by Naturhistoriska Museet, Goteborg, and DeutschesMeeresmuseum, Stralsund. The selection procedure aimed to include as many speciesas possible and to be as wide-ranging as possible in terms of geography and chrono-logy, and at the same time to avoid preserved specimens and specimens expected tocontain insufficient amounts of collagen.
A total of 15 different species was identified in the Danish collection (Table 1).Out of these it has been possible to date 12 by the radiocarbon method. In addition,10 nonspecified baleen whales were successfully radiocarbon dated, as these wereexpected to reveal important geological information.
The number of remains found in archaeological contexts makes up about 55% ofthe specimens and consists mainly of smaller toothed whales, especially the harbor
256 MARINE MAMMAL SCIENCE, VOL. 26, NO. 2, 2010
Table 1. Number of species identified and number of specimens dated in the Danishcollection of subfossil whale remains at ZMUC.
Number of specimensGeologically Archaeologically
Taxa 14C-dated dated dated Undated Total
L. albirostris 1 - 9 8 18D. delphis - - 2 - 2T. truncatus 1 - 15 18 34Stenella sp. - - 1 - 1Delphinidae Dolphins - - 3 2 5O. orca 4 1 16 40 61P. phocoena - - 49 14 63D. leucas 4 - 1 2 7H. ampullatus 1 - - 1 2P. macrocephalus 2 - 4 11 17B. acutorostrata 3 - 1 9 13B. physalus 2 - 1 7 10B. cf. musculus 1 - - 6 7Balaenoptera sp. 1 - 2 11 14Balaenopteridae Rorquals 1 - - - 1M. novaeangliae 2 - - 3 5E. glacialis 2 - - 3 5B. mysticetus 7 - 1 8 16B. mysticetus/E. glacialis 7 - - 10 17Cetacea ind. - 11 24 28 63Total 39 12 129 181 361
porpoise (P. phocoena). The harbor porpoise was omitted from the radiocarbon datingeffort because a relatively high number of archaeological datings already documentsits history in southern Scandinavia as shown by Sommer et al. (2008).
The remaining 45% of the whale bones were found in geological, that is, non-archaeological, contexts either in situ or redeposited. These bones mainly representlarge baleen whale or larger toothed whale species. The bones have been fished ordredged from the sea bottom or excavated in raised marine sediments includingold beach ridges or found redeposited in glacial and glaciofluvial deposits. Theexception to these generally abundant find scenarios is 13 adjoining lumbar andcaudal vertebrae of a B. mysticetus specimen and an almost complete front half of a finwhale (Balaenoptera physalus), both excavated in raised marine sediments in northernJylland (Table 2, 34 and 22). Normally subfossil whales were represented by a singleor a few bone elements only.
Identification
A safe identification of whale bones based on size and morphology completelydepends on the accessibility to a good reference collection of extant whale speciesskeletons. The ZMUC, houses a very large and diverse collection that has beensuccessfully used in this study. The subfossil remains, however, most often con-sist of rather fragmented vertebrae, skulls or ribs and more seldom of limb bones,mandibles or single teeth sometimes preventing identification to species level (see
AARIS-SØRENSEN ET AL.: SUBFOSSIL WHALE BONES 257
Tabl
e2.
List
ofan
alyz
edw
hale
rem
ains
and
the
resu
lts
ofth
era
dioc
arbo
nda
ting
(16
ofth
eda
ted
spec
imen
spl
uson
eun
date
dha
veal
sobe
enan
alyz
edfo
rtr
ace
elem
ents
;see
Tabl
e5)
.The
colu
mn
“Ele
men
tda
ted”
indi
cate
sw
hich
part
ofth
ew
hale
has
been
sam
pled
.The
colu
mn
“14C
met
hod”
indi
cate
sif
the
radi
ocar
bon
dati
ngha
sbe
enpe
rfor
med
byth
eco
nven
tion
alm
etho
dby
use
ofa
prop
orti
onal
coun
ter
(Con
v.)
orby
Acc
eler
ator
Mas
sSp
ectr
osco
py(A
MS)
.�13
Cis
the
stab
leis
otop
era
tio
13C
/12C
mea
sure
dag
ains
tV
ienn
aP
eeD
eeB
elem
nite
(VP
DB
),th
ein
tern
atio
nals
tand
ard
used
byal
lrad
ioca
rbon
labo
rato
ries
.The
cali
brat
edda
teis
give
nas
the
oute
rbo
unda
ries
cove
ring
the
inte
rval
±1
�.
Cal
ibra
ted
age
Ele
men
tZ
MU
CLo
cali
ty,
14C
14C
-age
�13C
BP
(Bef
ore
No.
Spec
ies
date
dfil
eco
unty
met
hod
Lab.
no.
BP
( �V
PD
B)
1950
)R
efer
ence
s
01B
.mys
tice
tus/
E.
glac
iali
sV
erte
bra
3/19
09Sp
arkæ
r,V
ibor
gC
onv.
K-7
098
Mod
ern
−15.
40
02B
.acu
toro
stra
taSk
ull,
occi
pita
lpa
rt22
/195
1Fø
nsso
gn,O
dens
eC
onv.
K-7
100
Mod
ern
−14.
70
03B
.phy
salu
sC
ervi
cal
vert
ebra
19/1
939
Hed
egar
dene
,Vej
leC
onv.
K-7
101
Mod
ern
−13.
50
04O
.orc
aR
ib4/
1932
Græ
rup,
Rib
eC
onv.
K-6
799
25±
70−1
1.5
0–26
005
H.a
mpu
llat
usV
erte
bra
40a/
1968
Kol
ding
Fjor
d,V
ejle
Con
v.K
-679
850
±70
−13.
10–
260
06T.
trun
catu
sSk
ull
93/1
946
Hin
dsga
ul,O
dens
eC
onv.
K-6
765
110
±50
−11.
80–
260
07M
.nov
aean
glia
eV
erte
bra
111/
1943
Søbo
rgR
uin,
Fred
erik
sbor
gC
onv.
K-6
795
705
±75
−14.
656
0–72
0
08B
alae
nopt
era
sp.
Ver
tebr
a32
5/19
82P
ouls
ker,
Bor
nhol
mC
onv.
K-6
804
1015
±70
−14.
880
0–1,
050
09M
ysti
ceti
Cer
vica
lve
rteb
ra28
/199
9D
anis
che
Wie
k,G
reif
swal
der
Bod
den,
Ger
man
y
Con
v.K
-709
511
10±
70−1
1.8
930–
1,12
0
10B
.mys
tice
tus/
E.g
laci
alis
Man
dibl
e37
/189
7Lø
nstr
up,H
jørr
ing
Con
v.K
-709
714
00±
75−1
5.5
1,27
0–1,
390
11B
.mys
tice
tus
Rad
ius
103/
1946
Stra
ndby
,Hjø
rrin
gC
onv.
K-6
820
1500
±65
−13.
51,
310—
1,51
012
B.m
ysti
cetu
s/E
.gla
cial
isLu
mba
rve
rteb
ra29
/199
9Z
ings
t,St
rals
und,
Ger
man
yC
onv.
K-7
096
1660
±65
−16.
21,
420–
1,69
0
13P.
mac
roce
phal
usH
umer
us17
/199
7Le
rbæ
kM
ark,
Hjø
rrin
gC
onv.
K-6
828
2560
±80
−14.
02,
490–
2,76
0
Con
tinu
ed
258 MARINE MAMMAL SCIENCE, VOL. 26, NO. 2, 2010Ta
ble2
.(C
onti
nued
)
Cal
ibra
ted
age
Ele
men
tZ
MU
CLo
cali
ty,
14C
14C
-age
�13C
BP
(Bef
ore
No.
Spec
ies
date
dfil
eco
unty
met
hod
Lab.
no.
BP
(�V
PD
B)
1950
)R
efer
ence
s
14O
.orc
aV
erte
bra
22/1
997
Ban
ners
lund
mar
k,H
jørr
ing
Con
v.K
-682
528
10±
75−1
2.4
2,79
0–3,
000
15B
.phy
salu
sSk
ull
1/19
21V
æng
esø,
Ran
ders
Con
v.K
-566
128
90±
80−1
5.1
2,89
0—3,
160
16P.
mac
roce
phal
usTo
oth
12/1
872
Læsø
,Kli
tgar
dN
.for
Byr
um,H
jørr
ing
Con
v.K
-257
629
20±
85−1
1.4
2,96
0–3,
210
(Han
sen
1977
)17
O.o
rca
Skul
l10
/194
1V
ust,
Thi
sted
Con
v.K
-676
632
50±
95−1
1.4
3,38
0–3,
580
18B
.cf.
acut
oros
trat
aSk
ull,
squa
mou
spa
rt
56/1
981
Nor
dby
Hed
e,Sa
msø
,H
olbæ
kC
onv.
K-6
770
3540
±90
-15.
43,
700–
3,96
0
19B
.mys
tice
tus
Skul
l66
/200
0R
orvi
k,B
ohus
Lan,
Swed
enA
MS
AA
R-7
984
4210
±80
−15.
724,
620–
4,85
0
20B
.mys
tice
tus
Skul
l,oc
cipi
tal
part
27/1
999
Lyng
by,T
hist
edC
onv.
K-7
094
4250
±90
−16.
14,
620–
4,96
0
21B
.mys
tice
tus/
E.g
laci
alis
Rib
1/19
68B
alle
rum
,Thi
sted
Con
v.K
-603
646
10±
85−1
4.4
5,07
0–5,
570
22B
.phy
salu
sV
erte
bra
12/1
988
Ast
ed,V
ibor
gC
onv.
K-5
998
5060
±95
−13.
85,
670–
5,91
0(S
tens
trop
1994
)23
B.a
cuto
rost
rata
Ver
tebr
a12
7/19
64H
olm
eeng
eI,
Ran
ders
Con
v.K
-680
351
30±
100
−14.
75,
740–
5,99
0
24B
alae
nopt
era
cf.
mus
culu
sSk
ull,
occi
pita
lpa
rt5/
1964
Tast
umSø
,Vib
org
Con
v.K
-679
751
60±
90−1
4.4
5,75
0–6,
000
25E
.gla
cial
isLi
mb
bone
71/1
965
Bal
leru
m,T
hist
edC
onv.
K-6
213
6410
±11
0−1
6.9
7,18
0–7,
430
26L
.alb
iros
tris
Skul
l5/
1896
Gni
ben,
Sjæ
llan
dsO
dde,
Hol
bæk
Con
v.K
-676
765
30±
110
−13.
07,
330–
7,56
0
27O
.orc
aSk
ull,
occi
pita
lpa
rt29
/198
3Ly
stru
pE
nge,
Arh
usC
onv.
K-6
800
6800
±11
5−1
1.8
7,53
0–7,
780
28D
.leu
cas
Ver
tebr
a60
/200
0P
ilga
rden
,Boh
usla
n,Sw
eden
AM
SA
AR
-797
898
80±
110
−13.
3911
,190
–11,
600
AARIS-SØRENSEN ET AL.: SUBFOSSIL WHALE BONES 259
29B
.mys
tice
tus
Ver
tebr
a65
/200
0Fr
andt
orps
gata
n,G
oteb
org,
Swed
enA
MS
AA
R-7
983
10,2
10±
80−1
6.05
11,7
60–1
2,07
0
30B
.mys
tice
tus
Pha
lanx
64/2
000
Bra
cke
smas
tuge
omra
de,
Got
ebor
g,Sw
eden
AM
SA
AR
-798
210
,915
±80
−15.
5112
,840
–12,
920
31D
.leu
cas
Cer
vica
lve
rteb
ra62
/200
0H
ulta
tege
lbru
k,V
aste
rgot
land
,Sw
eden
AM
SA
AR
-798
011
,090
±80
−12.
2412
,930
–13,
080
32B
.mys
tice
tus/
E.g
laci
alis
Rib
33/1
997
Em
mer
sbæ
k,H
jørr
ing
Con
v.K
-682
311
,320
±16
5−1
3.7
13,0
60–1
3,35
0
33B
.mys
tice
tus
Skul
l,oc
cipi
tal
part
19/0
000
Rah
olte
,Hjø
rrin
gC
onv.
K-7
108
11,5
90±
115
−13.
913
,310
–13,
570
34B
.mys
tice
tus
Ver
tebr
a11
/185
9B
ovbæ
k,H
jørr
ing
Con
v.K
-679
611
,780
±17
0−1
4.3
13,4
40–1
3,79
035
Bm
ysti
cetu
sR
ib20
/000
0B
orgb
akke
,Hjø
rrin
gC
onv.
K-7
109
11,9
60±
115
−13.
913
,720
–13,
950
36B
.mys
tice
tus
Rib
63/2
000
Tanu
msh
ede,
Boh
usLa
n,Sw
eden
AM
SA
AR
-798
111
,980
±90
−14.
8313
,750
–13,
940
37D
.leu
cas
Cau
dal
vert
ebra
176/
1980
Fred
erik
shav
nV
andv
ærk
,H
jørr
ing
Con
v.K
-710
412
,000
±17
0−1
2.3
13,6
90–1
4,06
0
38M
.cf.
nova
eang
liae
Rib
18/1
997
Ves
ter
Tvæ
rste
dSk
ole,
Hjø
rrin
gC
onv.
K-6
821
12,1
00±
185
−14.
713
,730
–14,
200
39B
.mys
tice
tus
Rib
27/1
953
Dro
nnin
glun
d,H
jørr
ing
Con
v.K
-710
512
,540
±15
5−1
5.1
14,3
40–1
4,95
0
40D
.leu
cas
Ver
tebr
a61
/200
0Sa
hlgr
ensk
aSj
ukhu
set,
Got
ebor
g,Sw
eden
AM
SA
AR
-797
912
,580
±90
−11.
8214
,580
–15,
020
41B
.mys
tice
tus/
E.g
laci
alis
Ver
tebr
a40
/199
6K
arup
,Hjø
rrin
gC
onv.
K-7
106
12,7
10±
185
−15.
114
,600
–15,
280
42B
.mys
tice
tus/
E.g
laci
alis
Ver
tebr
a31
/199
7H
irts
hals
,Hjø
rrin
gC
onv.
K-6
824
12,7
80±
215
−13.
514
,670
–15,
420
Con
tinu
ed
260 MARINE MAMMAL SCIENCE, VOL. 26, NO. 2, 2010
Tabl
e2.
(Con
tinu
ed)
Cal
ibra
ted
age
Ele
men
tZ
MU
CLo
cali
ty,
14C
14C
-age
�13C
BP
(Bef
ore
No.
Spec
ies
date
dfil
eco
unty
met
hod
Lab.
no.
BP
(�V
PD
B)
1950
)R
efer
ence
s
43B
.mys
tice
tus/
E.g
laci
alis
Ver
tebr
a26
0/19
80G
ølst
rup
Tegl
værk
sgra
v,H
jørr
ing
Con
v.K
-680
212
,950
±19
5−1
5.0
15,0
20–1
5,60
0
44B
.mys
tice
tus/
E.g
laci
alis
Ver
tebr
a26
1/19
80Sø
nder
Vra
,Hjø
rrin
gC
onv.
K-6
801
13,6
70±
205
−14.
915
,940
–16,
590
45B
.mys
tice
tus
Rib
21/0
000
Rav
nsho
lt,H
jørr
ing
Con
v.K
-711
014
,110
±21
5−1
4.1
16,4
70–1
7,20
046
E.g
laci
alis
Ver
tebr
a1/
1934
Stau
rby
Skov
,Ode
nse
Con
v.K
-676
817
,830
±59
0−1
7.9
20,4
70–2
1,92
047
Bal
aeno
pter
idae
Rib
16/1
928
Kro
gstr
upso
gn,
Fred
erik
sbor
gC
onv.
K-7
099
28,1
40±
1280
−15.
5-
48B
.acu
toro
stra
taC
auda
lve
rteb
ra32
/195
6Li
ndho
lm,N
ørre
Sund
by,A
lbor
gC
onv.
K-6
771
29,3
20±
1240
−13.
0-
49D
.leu
cas
Cau
dal
vert
ebra
41/1
952
Ves
ter
Neb
el,V
ejle
Con
v.K
-710
232
,220
±148
0−1
3.7
-
50D
.leu
cas
Cau
dal
vert
ebra
19/1
960
Nyh
olm
smar
k,H
jørr
ing
Con
v.K
-710
733
,140
±146
0−1
3.4
-
51D
.leu
cas
Cau
dal
vert
ebra
3/19
02A
sbak
ken,
Hjø
rrin
gC
onv.
K-7
103
>38
,320
−11.
1-
AARIS-SØRENSEN ET AL.: SUBFOSSIL WHALE BONES 261
Figure 1. Calibrated radiocarbon dates. The calibrations have been performed using theOXCAL calibration program and the 2004 IntCal04 curves.
Table 1). About one-third of the whale specimens in the ZMUC collection couldnot be identified to species level. The problems with the identification are espe-cially connected to the distinction between the two right whale species (makingup one-third of the dated specimens) and between the different rorquals (GenusBalaenoptera). Distinguishing between the two monodontid species (D. leucas and
262 MARINE MAMMAL SCIENCE, VOL. 26, NO. 2, 2010
Monodon monoceros) also presented some difficulty depending on the available boneelements. However, one of the seven specimens of Monodontidae found in Den-mark, could be identified with certainty to D.leucas from a cervical vertebra (at-las) (Winge 1899). Besides, several complete or almost complete skeletons ofD. leucas have been found on the Swedish west coast (Lepiksaar 1966, Freden 1984).So far no subfossil remains of M. monoceros have been identified in southern Scandi-navia. Therefore, the other six Danish specimens (all caudal vertebrae) have similarlybeen assigned to D. leucas.
Radiocarbon Dating
In the present work, we undertook the radiocarbon dating of many new specimensof whales found in either archaeological or geological contexts. Out of the 41 Danishand 2 German whale bone samples, 39 were dated successfully at the CopenhagenRadiocarbon Laboratory utilizing a conventional proportional counter (Rasmussen2000). The remaining four samples contained insufficient amounts of collagen. Theseven Swedish specimens were dated by the Aarhus AMS radiocarbon laboratory forour work. By adding another five samples dated earlier, a grand total of 51 dates arelisted in Table 2.
The stable isotope ratio, �13C, was measured on all 51 samples, and the dates werecorrected for isotopic fractionation referring them to the marine value of 0� VPDB.This corresponds to subtracting a reservoir correction of 405 yr from the standardradiocarbon age, which is referred to �13C = −25� VPDB.
Three of the samples were dated to be modern, and one older than the datinglimit. The dates that could be calibrated were calibrated with the OXCAL programusing the Reimer et al. (2004) curves. The calibrated date intervals at ±1 SD arelisted in Table 2, and shown in Figure 1.
Trace Element Determination
Seventeen samples were subjected to trace element analysis. This was done usingInstrumental Neutron Activation Analysis (INAA) on ca. 500-mg samples of bonetissue. In order to reduce the effect of contamination present on the surface of thewhale bones, an exterior layer of ca. 1 mm of the bone was removed with a drill orwith a scalpel prior to sampling. The samples were irradiated at the now disassembledheavy water reactor DR3 at the National Laboratory at Risø, Denmark, with a typicalneutron flux of 1013 cm−2s−1 and an irradiation time of 4 h. Standard samples aswell as flux monitors were included in the irradiation. The activated samples weresubjected to three counts on a high purity GeLi-detector at the Geological Institute,University of Copenhagen. All samples were analyzed only once. The resultingconcentrations are listed in Table 3. The uncertainties are mostly in the range of10%–15%.
RESULTS AND DISCUSSION
The Subfossil and Recent Cetacean Fauna
The results of the radiocarbon dating are compiled in Table 2 together with basicinformation about each specimen. The chronological spread of the dates is visualized
AARIS-SØRENSEN ET AL.: SUBFOSSIL WHALE BONES 263
Tabl
e3.
Chr
onol
ogic
aldi
stri
buti
onof
25w
hale
spec
ies
inth
eD
anis
h/so
uthe
rnSc
andi
navi
anar
ea.
a:14
C-d
ates
this
stud
y;b:
14C
-dat
e,K
yvik
,H
alla
nd,S
wed
en(F
rede
n19
84);
c:ge
olog
ical
lyda
ted,
c1:U
ddev
alla
,Boh
usla
n,Sw
eden
(Lep
iksa
ar19
66,1
986,
Fred
en19
84),
c2:O
tter
o,B
ohus
lan,
Swed
en(L
epik
saar
1966
);d:
arch
aeol
ogic
ally
date
d,d1
:se
vera
llo
cali
ties
,Z
MU
Cfil
es,
d2:
Tybr
ind
Vig
,Fy
n(T
roll
e-La
ssen
1985
);e:
(Kin
ze20
07);
f:(K
inze
2006
a);
g:pe
rson
alco
mm
unic
atio
nfr
omU
noSv
enss
on,
Got
ebor
gsN
atur
hist
oris
kaM
useu
m,
Apr
il20
07;
h:(L
epik
saar
1966
).∗ M
arin
een
viro
nmen
tba
sed
onm
ollu
skst
udie
saf
ter
Pet
erse
n(2
004)
.
Old
erY
oldi
aSe
aY
oung
erY
oldi
aSe
aar
ctic
-sub
arct
ic∗
arct
ic-s
ubar
ctic
-bor
eal∗
Litt
orin
a/Ta
pes
Sea
Late
Mid
dle
Wei
chse
lian
-LG
MLG
M-P
lei/
Hol
boun
dary
bore
al-l
ucit
ania
n∗
Spec
ies
ca.3
3–21
cal.
kyr
BP
ca.2
1–11
.7ca
l.ky
rB
Pca
.8.0
–0ca
l.ky
rB
PR
ecen
tfa
una
L.a
lbir
ostr
isX
c1X
aX
ena
tive
L.a
cutu
sX
eco
mm
onD
.del
phis
Xd1
Xe
occa
sion
also
uthe
rnvi
sito
rT.
trun
catu
sX
aX
eoc
casi
onal
sout
hern
visi
tor
Sten
ella
sp.
Xd2
Xe
occa
sion
also
uthe
rnvi
sito
rG
lobi
ceph
ala
mel
asX
eco
mm
onP.
cras
side
nsX
era
revi
sito
rF
eres
aat
tenu
ata
Xf
rare
visi
tor
O.o
rca
Xc2
Xa
Xe
com
mon
Gra
mpu
sgri
seus
Xe
rare
visi
tor
P.ph
ocoe
naX
bX
d1X
ena
tive
D.l
euca
sX
aX
aX
d1X
era
revi
sito
rM
.mon
ocer
osX
gra
revi
sito
rM
esop
lodo
nbi
dens
Xe
rare
visi
tor
H.a
mpu
llat
usX
c1X
aX
era
revi
sito
rZ
.cav
iros
tris
Xh
rare
visi
tor
P.m
acro
ceph
alus
Xa
Xe
erra
tic
and
peri
odic
ally
freq
uent
stra
ggle
rB
.acu
toro
stra
taX
aX
aX
eco
mm
onB
alae
nopt
era
bryd
eiX
era
revi
sito
rB
alae
nopt
era
bore
alis
Xe
rare
visi
tor
B.p
hysa
lus
Xa
Xe
peri
odic
ally
freq
uent
B.c
f.m
uscu
lus
Xa
Xe
rare
visi
tor
M.n
ovae
angl
iae
Xa
Xa
Xe
rare
visi
tor
E.g
laci
alis
Xa
Xa
Xe
rare
visi
tor
B.m
ysti
cetu
sX
aX
a
264 MARINE MAMMAL SCIENCE, VOL. 26, NO. 2, 2010
Kattegat
Skagerrak
Baltic Sea
resundBeltSea
Nort
h S
ea
Sjælland
Fyn
Jylland
Bohuslän
Västergötland
Halland
Bornholm
DENMARK
GERMANY
SWEDEN
44
42
3210
39 16
34
50
51
45
4341
38
48
18
17
3
2
1
4
5
6
7
8
15
21
26
20
22
23
24
27
46
47
49
19
36
29, 30, 40
28
31
11,13, 14, 33, 35,
12
9
37
25
Figure 2. Map of southern Scandinavia showing the distribution of the finds. Numbersrefer to specimen numbers in Table 2.
in Figure 1 and the geographical distribution of the dated whales is shown inFigure 2.
The dates fall into three well-defined groups. The oldest group (five dates) rangefrom ca. 33.0–21.0 cal. kyr BP corresponding to a period during the Weichselianwith prevailing interstadial conditions beginning in the late Middle Weichselianand ending with the Last Glacial Maximum (LGM) (Fig. 3a). The next group(18 dates) covers the deglaciation period between the LGM and the Pleis-tocene/Holocene boundary, ca. 17.0–11.7 cal. kyr BP (Fig. 3c). Finally, the youngestgroup (24 dates) range between ca. 8.0 cal. kyr BP and the present time correspondingto the time after the Early Atlantic transgression and the formation of the Danishislands (Fig. 4). In terms of climate and marine environments the three periods
AARIS-SØRENSEN ET AL.: SUBFOSSIL WHALE BONES 265
Figure 3. Palaeogeographical reconstructions of southern Scandinavia during the late Mid-dle Weichselian and the Late Weichselian. (a) interstadial, ca. 33–31 cal. kyr BP. (b) the LastGlacial Maximum ca. 23–21 cal. kyr BP. (c) final deglaciation, ca. 16–14.5 cal. kyr BP. In aand c: A palaeo-Kattegat-Skagerrak with drift ice and icebergs south of the Norwegian icefront is connected with a Baltic Ice Lake. Gray shaded land areas are dominated by dead-ice.(After Houmark-Nielsen et al. 2005). This figure is available in color online.
correspond to the arctic-subarctic Older Yoldia Sea, the arctic-subarctic-borealYounger Yoldia Sea and the boreal-lucitanian Littorina/Tapes Sea.
A majority of the dated specimens belonging to the two oldest groups, includ-ing a single nonfinite date of >38 kyr BP (K-7103), were found in northernJylland and Bohuslan on the Swedish west coast. This is in agreement with theglacial history and palaeogeography as outlined by recent geological studies (see e.g.,
266 MARINE MAMMAL SCIENCE, VOL. 26, NO. 2, 2010
Figure 3. (Continued)
Houmark-Nielsen and Kjær 2003, Houmark-Nielsen et al. 2005), which impliesa geographical distribution of the Older and Younger Yoldia Sea corresponding tothe present Skagerrak and Northern Kattegat (Fig. 3a, c) and occasionally reachingas far south as Northern Sjælland (Bahnson et al. 1974, Petersen and Buch 1974).In accordance with this the members of the youngest group have been found muchmore dispersed and further south following the later emerging coast lines along theinner Danish waters (Fig. 4).
The geologically and archaeologically dated specimens listed in Table 1support the results obtained through the radiocarbon dating. The re-mains of Delphinus delphis, Stenella sp., and P. phocoena have, however, beensolely archaeologically dated which place the two dolphins within the Late
AARIS-SØRENSEN ET AL.: SUBFOSSIL WHALE BONES 267
Figure 3. (Continued)
Atlantic–Early Subboreal (Late Ertebølle–Early Neolithic Cultures) and the har-bor porpoise between the Early Atlantic (Kongemose Culture) and the present time.The latter is in agreement with the conclusion by Sommer et al. (2008) that placesthe first immigration of the harbor porpoise into the Baltic Sea at 9.0–7.5 cal. kyrBP. A much earlier occurrence is, however, known from the Kattegat area where aspecimen found on the Swedish west coast at Kyvik, Halland (Lepiksaar 1966) hasbeen radiocarbon dated to ca. 12.9–12.0 cal. kyr BP (Freden 1984) (Table 3). Thisindicates the presence of the harbor porpoise in the Skagerrak/Kattegat area since thelate glacial followed by an expansion into the inner Danish waters and the Baltic Seain the wake of the early Atlantic transgression. Table 3 summarizes the chronolog-ical distribution of the 15 whale species in the Danish/southern Scandinavian area
268 MARINE MAMMAL SCIENCE, VOL. 26, NO. 2, 2010
Figure 4. Palaeogeographical map of southern Scandinavia around 6.5 cal. kyr BP. Atlantictransgressions have created the Danish islands and inner waters (drawing by Knud Rosenlund).This figure is available in color online.
based primarily on this study, but supplemented with earlier radiocarbon dated andarchaeologically dated specimens available in the ZMUC files and the literature. Incomparison with the recent cetacean fauna the following differences and similaritiesshould be noted.
The waters around the present Denmark include part of the North Sea, Skagerrak,Kattegat, the Belt Sea, and the Øresund as well as the westernmost Baltic properand span in depth from 0 to several hundred meters.
Twenty-two whale species have been documented from the recent Danish watersalone, and two additional species are known from adjacent Swedish waters, that is,
AARIS-SØRENSEN ET AL.: SUBFOSSIL WHALE BONES 269
Cuvier’s beaked whale (Ziphius cavirostris) and narwhale (Monodon monoceros), seeTable 3. Among these there are two native species, the harbor porpoise (P. pho-coena) and the white-beaked dolphin (L.albirostris); four common species, the Atlanticwhite-sided dolphin (Lagenorhynchus acutus), the long-finned pilot whale (Globicehalamelas), the killer whale (O. orca), and the common minke whale (Balaenoptera acu-torostrata); and 18 more or less frequently occurring stragglers from waters eitherdiffering in temperature, in water depth, or both. Their occurrence is linked tohydrographical phenomena such as saltwater intrusions and fluctuations in surfacewater temperature (Kinze et al. 2001).
The two native species and two of the common species (killer whale and minkewhale) are also among the most frequently found subfossil species documenting theirlong presence in southern Scandinavia (Table 1). The two other common species, theAtlantic white-sided dolphin and the long-finned pilot whale are, however, totallyabsent in the subfossil record. This is probably due to the fact that they are bothoffshore species normally found in deeper waters in the Danish sector of the North Seaand Skagerrak. A low number of strandings along the coast of the inner Danish watersin the past should therefore be expected. Only two incidents of mass strandings areknown from the inner Danish waters during the last century, one of the white-sideddolphin in Roskilde Fjord in 1942 and one of the pilot whale in Vejle Fjord in 1954(Kinze 1995).
Among the baleen whales the list comprises four species that do possess the capacityto dwell in coastal waters. These are, besides the already mentioned common minkewhale (B. acutorostrata), the fin whale (B. physalus), the humpback whale (Megapteranovaeangliae), and the North Atlantic right whale (E. glacialis). The subfossil recordsuggests that the latter has visited the Danish waters ever since the Weichselianand probably have had a more common occurrence before the severe exploitationduring historical times. The other right whale, the bowhead (B. mysticetus), has alsoalmost been wiped out by commercial whaling in historical times and has neverbeen recorded in the local national stranding lists (Kinze 1995, 2006b, Kinze et al.1998, 2001). On the other hand it is well represented in the subfossil record wherethe 16 specimens from Danish collections can be supplemented with 30 finds fromthe Swedish west coast (Freden 1984) of which four have been included in thepresent dating program. The majority of the dates fall within the arctic-subarcticlate Glacial period that is in accordance with the modern biology of this circum-arcticwhale which is seldom sighted south of 45◦N. A few young dates ranging betweenca. 1.5 and 5.0 cal. kyr BP nevertheless indicate that the bowhead was an occasionalvisitor to southern Scandinavian waters during post-glacial times prior to modernexploitation.
Another witness of the full glacial/late glacial environment is the beluga whale(D. leucas). Today this arctic coastal species occasionally conducts extralimited mi-grations into more southerly waters including the inner Danish waters (Kinze 2007).This is in accordance with the subfossil record that only reveals a single Danish findfrom the boreal-lucitanian Littorina/Tapes Sea but several Swedish and Danish findsfrom the arctic-subarctic Yoldia Sea.
Three warm temperate to tropical dolphin species (D. delphis, Stenella sp., andT. truncatus) indicate much warmer sea water temperatures during earlier peri-ods. They have all been found in archaeological contexts and hereby dated tothe Late Atlantic–Early Subboreal period corresponding to the postglacial climaticoptimum.
270 MARINE MAMMAL SCIENCE, VOL. 26, NO. 2, 2010
Finally it should be noted that the northern bottlenose (Hyperoodon ampullatus)as well as the sperm whale (P. macrocephalus) both are strictly oceanic species thatget beached accidentally either on shallow tidal flats or in coastal areas of complexhydrography. This seems to have happened in the past as well.
Trace Element Concentrations
The trace element concentrations determined in this work are listed in Table 4.The first question to be raised is whether bone diagenesis has occurred on a largescale. If diagenesis has occurred in a systematic way over a scale of hundreds orthousands of years it is likely that it would lead to progressive loss of Ca withincreasing age. However, no correlation is seen between Ca and age, the correlationcoefficient is r2 = 0.0013. Another conceivable way diagenesis might be manifestedis by progressive loss of collagen with increasing age. No correlation is seen in thiscase either, the correlation coefficient is calculated to be r2 = 0.0216. Accordingto these two criteria we see no signs of long-term diagenesis. However, it is stillpossible that diagenesis has occurred on a shorter time scale not reflected in thesecriteria. If this has been the case it is likely that the diagenetic processes encompasssingle elements related to the marine environment, for example, as demonstrated forAs in Mesolithic human and animal bones buried in an area later covered by the sea(Rasmussen et al. 2009). Guided by the findings of Rasmussen et al. (2009) we findit likely that a variation in the As concentration in the bones could be ascribed toprocesses involving decomposition of kelp or other marine plants with a high contentof As, and therefore variations in As is not considered worthwhile to look at in thisconnection.
Trueman et al. (2004) showed massive diagenesis in bones deposited on the surfaceof the savannah in Africa, but these alterations were ascribed to the osmotic raise ofground water from the soil, through the bone and followed by evaporation from thebone, thus leading to an ever increasing concentration of the trace elements in thebone. Trueman et al. (2004) also showed collagen to disappear within decades afterdeposition on the surface. However, our scenario is radically different, as the whalebones were deposited on the seafloor in the somewhat anoxic, cold conditions of theinner Danish waters. We observed only four cases out of 43 where complete collagenremoval have taken place. So there is no reason to think that the results of Truemanet al. (2004) will apply to our samples. We have also shown that Hg is probably notmobilized in human bones buried in Danish soil (Rasmussen et al. 2008) lendingcredence to the assumption that bones are not always subjected to diagenesis.
Even if it could be assumed that diagenesis has not occurred to any large extent, aninvasion of terrigenic clastic material has almost certainly occurred. Small sedimentparticles from the surrounding sea bed have undoubtedly invaded the pore spacesin the porous and less competent parts of the bone. Even though great care hasbeen taken to decontaminate the surface of the bone during sampling, it is mostlikely that soil particles at least to a minor degree are part of the samples analyzed.Examples of elements that must be considered to originate mainly from invasion ofterrigenic clastic material are the REE (Rare Earth Elements), eight of which arelisted in Table 4, as well as Sc, Ta, and Th. Consequently, we have also eliminatedthese elements from use in this study.
Even if diagenesis might occur for some elements, it is, however, also possiblethat the dietary habits are reflected in the distribution of some of the trace elements.
AARIS-SØRENSEN ET AL.: SUBFOSSIL WHALE BONES 271
Tabl
e4.
Trac
eel
emen
tco
ncen
trat
ions
.“nd
”m
eans
not
dete
rmin
ed.T
heco
llag
enco
nten
tis
dete
rmin
edby
wei
ghin
gth
eor
igin
albo
nesa
mpl
ean
dw
eigh
ing
the
drie
dco
llag
enpr
oduc
edpr
ior
tora
dioc
arbo
nda
ting
.The
unce
rtai
ntie
sof
the
INA
Are
sult
sar
ege
nera
lly
betw
een
10%
and
15%
.
14C
-age
Col
lage
nLa
bno
.n
Sam
ple
loca
tion
Kno
.B
P19
50E
rror
±1
��13
Cw
t%
KLR
-169
96
Hin
dsga
vlK
-676
511
050
−11.
815
.6K
LR-1
700
17V
ust
K-6
766
3250
95−1
1.4
10.8
KLR
-170
126
Sjæ
llan
dsO
dde,
K-6
767
6530
110
−13.
010
.6K
LR-1
702
46St
auer
byK
-676
817
,830
590
−17.
910
.5K
LR-1
703
ndA
albo
rgK
-676
9nd
ndnd
1.7
KLR
-170
418
Nor
dby
hede
,Sam
søK
-677
035
4090
−15.
418
.5K
LR-1
705
48Li
ndho
lm,A
albo
rgK
-677
129
,320
1240
−13.
09.
0K
LR-1
706
7Sø
borg
ruin
K-6
795
705
75−1
4.6
5.1
KLR
-170
734
Bov
bæk,
Ugg
erby
,Hjø
rrin
gK
-679
611
,780
170
−14.
313
.5K
LR-1
708
24Ta
stum
sø,V
ibor
gK
-679
751
6090
−14.
44.
8K
LR-1
709
5K
oldi
ngfj
ord
K-6
798
5070
−13.
112
.3K
LR-1
710
4G
ræru
p,O
ksbø
lK
-679
925
70−1
1.5
11.2
KLR
-171
127
Lyst
rup
enge
,Ele
vso
gnK
-680
068
0011
5−1
1.8
11.5
KLR
-171
244
Sdr.
Vra
,Hjø
rrin
gK
-680
113
,670
205
−14.
99.
6K
LR-1
713
43G
ølst
rup
tegl
v.H
jørr
ing
K-6
802
12,9
5019
5−1
5.0
7.5
KLR
-171
423
Hol
me
enge
,Ran
ders
K-6
803
5130
100
−14.
715
.8K
LR-1
715
8P
ouls
ker,
Bor
nhol
mK
-680
410
1570
−14.
86.
5
Con
tinu
ed
272 MARINE MAMMAL SCIENCE, VOL. 26, NO. 2, 2010
Tabl
e4.
(Con
tinu
ed)
Na
Ca
ScC
rFe
Co
Zn
Lab
no.
�g/
g�
g/g
�g/
g�
g/g
�g/
g�
g/g
�g/
g
KLR
-169
95,
040
235,
000
0.11
0.48
231
0.08
306
KLR
-170
05,
490
197,
000
1.03
7.54
3,42
00.
8533
0K
LR-1
701
5,17
027
6,00
00.
341.
2337
10.
1728
6K
LR-1
702
4,14
021
9,00
01.
434.
4092
802.
9110
7K
LR-1
703
4,57
025
7,00
01.
2718
.10
33,5
0050
.50
169
KLR
-170
43,
600
265,
000
2.11
5.50
12,1
001.
4117
2K
LR-1
705
4,69
026
6,00
00.
483.
619,
540
0.47
87K
LR-1
706
4,61
020
0,00
02.
0610
.70
12,8
002.
9312
4K
LR-1
707
410
231,
000
0.05
0.10
1,19
00.
0213
KLR
-170
86,
280
208,
000
0.97
10.6
019
,000
0.71
70K
LR-1
709
8,37
019
6,00
01.
4310
.90
6,97
01.
7526
5K
LR-1
710
3,45
028
2,00
00.
02nd
108
0.07
481
KLR
-171
15,
710
231,
000
1.31
5.16
3,63
02.
1422
1K
LR-1
712
4,36
021
1,00
05.
848.
3114
,900
1.87
52K
LR-1
713
4,55
025
7,00
01.
254.
8517
,800
17.8
082
KLR
-171
433
825
2,00
00.
100.
403,
380
0.11
12K
LR-1
715
2,17
033
2,00
00.
241.
8887
50.
3715
8
AARIS-SØRENSEN ET AL.: SUBFOSSIL WHALE BONES 273
As
Br
Rb
SrSb
Cs
Ba
Lab
no.
�g/
g�
g/g
�g/
g�
g/g
�g/
g�
g/g
�g/
g
KLR
-169
91.
546.
10.
461
40.
090
nd77
KLR
-170
02.
3912
.812
.525
60.
231
0.29
119
KLR
-170
12.
669.
4nd
1,03
00.
290
nd25
2K
LR-1
702
13.8
033
.711
.641
80.
520
0.37
337
KLR
-170
321
7.00
15.7
9.2
1,26
03.
520
0.32
1,26
0K
LR-1
704
15.0
029
.3nd
883
1.19
0nd
279
KLR
-170
56.
911.
43.
879
40.
045
0.25
88K
LR-1
706
5.89
28.7
24.0
1,92
00.
356
0.67
362
KLR
-170
70.
170.
20.
138
0.00
3nd
12K
LR-1
708
18.7
07.
09.
463
30.
714
0.13
325
KLR
-170
96.
4661
.212
.690
00.
398
0.55
126
KLR
-171
0nd
16.8
nd23
10.
028
ndnd
KLR
-171
12.
476.
68.
180
10.
239
0.18
286
KLR
-171
216
.30
nd16
.350
90.
222
0.51
342
KLR
-171
320
7.00
5.5
10.0
533
0.58
80.
311,
460
KLR
-171
45.
611.
5nd
305
0.04
0nd
51K
LR-1
715
2.33
21.1
4.5
503
0.09
70.
1727
Con
tinu
ed
274 MARINE MAMMAL SCIENCE, VOL. 26, NO. 2, 2010
Tabl
e4.
(Con
tinu
ed)
LaC
eN
dSm
Eu
Tb
Yb
Lab
no.
�g/
g�
g/g
�g/
g�
g/g
�g/
g�
g/g
�g/
g
KLR
-169
91.
1nd
nd0.
130
0.03
40.
023
0.05
3K
LR-1
700
10.3
9.1
6.4
0.84
50.
235
0.21
00.
839
KLR
-170
113
.03.
54.
0nd
0.05
30.
064
0.31
2K
LR-1
702
25.1
12.9
9.5
0.69
60.
270
0.24
11.
020
KLR
-170
318
8.0
49.5
36.0
nd0.
162
0.14
40.
498
KLR
-170
427
.842
.635
.75.
140
1.13
01.
070
2.92
0K
LR-1
705
1.8
2.4
2.3
0.21
80.
045
0.02
90.
136
KLR
-170
68.
217
.08.
41.
410
0.27
50.
220
0.88
4K
LR-1
707
0.3
0.5
0.2
0.06
00.
012
0.01
20.
066
KLR
-170
844
.613
.98.
50.
138
0.15
80.
140
0.86
6K
LR-1
709
13.4
10.7
5.0
0.70
30.
171
0.13
70.
286
KLR
-171
00.
6nd
ndnd
0.00
6nd
0.13
3K
LR-1
711
42.6
18.4
9.5
0.79
00.
263
0.25
91.
130
KLR
-171
230
.259
.529
.95.
340
1.10
01.
230
5.86
0K
LR-1
713
852.
050
.737
.2nd
0.11
80.
104
0.47
4K
LR-1
714
7.5
11.8
4.7
0.56
50.
113
0.11
40.
323
KLR
-171
50.
8nd
nd0.
146
0.02
60.
016
nd
AARIS-SØRENSEN ET AL.: SUBFOSSIL WHALE BONES 275
LuH
fTa
Th
Lab
no.
�g/
g�
g/g
�g/
g�
g/g
KLR
-169
9nd
ndnd
0.04
0K
LR-1
700
0.11
41.
860
0.10
900.
880
KLR
-170
10.
058
0.05
6nd
0.10
0K
LR-1
702
0.14
70.
585
0.07
410.
755
KLR
-170
30.
074
2.26
00.
1690
0.96
9K
LR-1
704
0.42
00.
229
nd1.
610
KLR
-170
5nd
0.28
70.
0231
0.39
3K
LR-1
706
0.11
12.
040
0.14
301.
770
KLR
-170
70.
009
0.00
90.
0019
0.01
7K
LR-1
708
0.12
23.
980
0.16
100.
791
KLR
-170
90.
028
1.40
00.
1140
0.97
1K
LR-1
710
nd0.
318
nd0.
047
KLR
-171
10.
138
1.98
00.
0766
0.99
2K
LR-1
712
0.95
13.
550
0.16
801.
620
KLR
-171
30.
066
0.59
70.
0751
0.62
9K
LR-1
714
0.04
40.
006
nd0.
076
KLR
-171
5nd
0.22
30.
0253
0.22
6
276 MARINE MAMMAL SCIENCE, VOL. 26, NO. 2, 2010
Table 5. Preferred diets of the specimens subjected to trace element analysis. The numbersin the first column are identical to specimen numbers in the first column of Table 2.
Diet knownNo. Species ID Species group literature
7 M. novaeangliae Søborg Ruin,Frederiksborg
Krill/fish
8 Balaenoptera sp. Poulsker,Bornholm
18 B. acutorostrata Nordby Hede,Samsø
Krill/fish
23 B. acutorostrata Holmeenge I,Randers
Krill/fish
48 B. acutorostrata Lindholm,Aalborg
Krill/fish
24 B. musculus, Tastum Sø, Viborg Krill46 E. glacialis, Staurby Skov,
OdenseCopepods
43 B. mysticetus/ E.glacialis,Gølstrup TeglværksgravHjørring
Copepods
34 B. mysticetus, Bovbæk, Hjørring Copepods44 B. mysticetus, Sønder Vra,
HjørringCopepods
K6769 B. mysticetus, Aalborg Copepods6 T. truncatus, Hindsgaul, Odense Fish/squid26 L. albirostris, Gnibben, Sjællands
OddeFish/squid
4 O. orca, Grærup, Ribe Fish/mammals17 O. orca, Vust, Thisted Fish/mammals27 O. orca, Kystrup Enge, Arhus Fish/mammals5 H. ampullatus, Kolding Fjord Squid
In order to investigate this, we have listed the major food preference for the whalespecies in this study in Table 5.
The food preferences do indeed show up, particularly in Fe, Cr, and Zn. Based onZn vs. Fe plot (Fig. 5) there is a clear distinction between crayfish eaters on one handand fish and squid eaters on the other. A similar if not quite so clear division is seenfor Cr vs. Fe (Fig. 6). This rather clear distinction or grouping could be caused by adifference in the amount of, or the chemistry of, the invasive sedimentary particles,as described above. If the difference arose this way, it can either be due to a differencein bone porosity upon degradation of the organic parts of the different species, or itcould be caused by a preference of special types of seabed where one species chooses todie. As an alternative explanation, it is possible that genetic differences between thewhale species are responsible for both a different dietary preference and a differentability to incorporate certain trace elements into the skeletal organ.
Finally, it is possible that the trace element composition pattern is transferred in auniform way from the food items to the bones of the whales. In this simple scenario thetrace element chemistry of the bones reflects whatever the whales forage on. With thepresent data, we are not able to distinguish between these alternatives. In addition,
AARIS-SØRENSEN ET AL.: SUBFOSSIL WHALE BONES 277
Figure 5. Zink vs. Fe, both in ppm (�g/g). The color code for the food preference is: red(or horizontal bars) = crayfish; green (or dots) = fish; yellow (or vertical bars) = squid; blue(or filled black) = mammals. This figure is available in color online.
we have not shown where the trace elements are situated—in the bone carbonatefraction, in the organic fractions, the nervous tissue or the fat, or in terrigenic clasticsediment grains.
Irrespective of the mechanism responsible the distribution of the trace elementsFe, Cr, and Zn seems none the less to be a way to distinguish between crayfish eatersfrom fish/squid eaters. It should therefore be possible to use these divisions in theelemental cross plots as a help in determining species. As an example, it can be seenfrom Figures 5 and 6 that specimen number 8 from Poulsker, Bornholm, which bymorphology has been determined and registered in the collection as Balaenoptera sp.,is likely to have been a fish/squid eater. This challenges the identification and makesit worth considering whether this fragmented vertebral body in fact belongs to asperm whale.
Conclusions
Based on our investigations we draw the following conclusions:
1. The subfossil whale record of southern Scandinavia comprises at least 15different species. Except for the bowhead whale (B. mysticetus) they are alldocumented in the modern fauna as well.
2. The modern fauna counts 24 species of which only two, the harbor porpoise (P.phocoena) and the white-beaked dolphin (L. albirostris) are native species to thearea. Another four species, the Atlantic white-sided dolphin (L. acutus),the long-finned pilot whale (G. melas), the killer whale (O. orca) and thecommon minke whale (B. acutorostrata) are common in southern Scandin-avian waters while the rest are more or less frequently occurring stragglers.
278 MARINE MAMMAL SCIENCE, VOL. 26, NO. 2, 2010
Figure 6. Chromium vs. Fe, both in ppm (�g/g). The color code for the food preference is:red (or horizontal bars) = crayfish; green (or dots) = fish; yellow (or vertical bars) = squid;blue (or filled black) = mammals. This figure is available in color online.
3. The harbor porpoise, the white-beaked dolphin, the killer whale and thecommon minke whale are also among the most frequently found subfossilspecies while the two offshore species, the white-sided dolphin and the pilotwhale, are absent from the subfossil record as they probably stranded veryrarely in the inner Danish waters.
4. The two right whales, the bowhead (B. mysticetus) and the North Atlanticright whale (E. glacialis), seem to have been occasional visitors to the southernScandinavian waters prior to the severe exploitation during historical times.
5. The large-scale climatic events during the last glacial-interglacial cycle arereflected in the whale fauna. Two species, the north Atlantic/arctic-subarcticbeluga whale (D. leucas) and the minke whale (B. acutorostrata), were recordedfrom the time prior to the LGM, (>38.0–ca. 21.0 kyr BP), eight species fromthe deglaciation period between LGM and the Pleistocene/Holocene bound-ary (ca. 17.0–11.7 cal. kyr BP) including the circum-arctic bowhead whale(B. mysticetus) and fifteen species, including warm temperate/tropical dolphins,during the Holocene from ca. 8.0 cal. kyr BP and onward.
6. The whale remains belonging to the two oldest groups have all been foundin northern Jylland and Bohuslan on the Swedish west coast. This is in ac-cordance with the palaeogeographical settings presented by recent geologicalinvestigations and it seems that a southern Scandinavian whale fauna existedduring the Weichselian glaciation whenever interstadial conditions created apalaeo-Skagerrak-Kattegat (Older Yoldia Sea and Younger Yoldia Sea). Re-mains belonging to the youngest group have been found much more dispersedand further south in accordance with the creation of the inner Danish watersby the transgressing Littorina/Tapes Sea.
7. Trace element cross plots exhibit distinct differences between certain whalespecies. With cross plots of the element concentrations of Fe, Zn, and Cr, it
AARIS-SØRENSEN ET AL.: SUBFOSSIL WHALE BONES 279
is possible to distinguish crayfish eaters from fish/squid eaters. This can beused as a novel and independent method to aid the determination to speciesof whale remains.
ACKNOWLEDGMENTS
Knud Rosenlund and Jeppe Møhl are thanked for help during the collecting and samplingof the many specimens, the late Karen Skov Jensen for technical assistance in the radiocarbonlaboratory, and Raymond Gwozdz for performing the INAA analyses. We are grateful to thecurators, Uno Svensson at the Goteborgs Naturhistoriska Museum and Harald Benke at theDeutsches Meeresmuseum, Stralsund, for access to the whale collections of the two museums.
LITERATURE CITED
Aaris-Sørensen, K. 1998. Danmarks forhistoriske Dyreverden [The prehistoric fauna of Den-mark]. Gyldendal, Copenhagen, Denmark.
Aaris-Sørensen, K. 2007. Fra istid til nutid (with an English summary: Late and Post Glacialmammals in Denmark). Pages 312–321 in H. J. Baagøe and T. S. Jensen, eds. DanskPattedyr Atlas [Atlas of Danish mammals]. Gyldendal, Copenhagen, Denmark.
Aurivillius, C. W. S. 1888. Der Wal Swedenborg’s nach einem Funde im Diluvium Schwedensbeurteilt. Kungliga Svenska Vetenskapsakademins Handlingar 23(1). 58 pp.
Bahnson, H., K. S. Petersen, P. B. Konradi and K. L. Knudsen 1974. Stratigraphy of Qua-ternary deposits in the Skærumhede II boring: Lithology, molluscs and foraminifera.Danmarks geologiske Undersøgelser, Arbog 1973:27–62.
Degerbøl, M. 1933. Danmarks Pattedyr i Fortiden i sammenligning med recente Former[The prehistoric mammals of Denmark compared with recent forms]. VidenskabeligeMeddelelser fra Dansk naturhistorisk Forening 96:357–641.
Freden, C. 1975. Subfossil finds of arctic whales and seals in Sweden. Sveriges GeologiskaUndersokning C 710. 62 pp.
Freden, C. 1984. Faunahistoriska notiser om nagra av Naturhistoriska Museets dateradesubfossila fynd [Faunal history notes on some of the dated subfossil finds in the NaturalHistory Museum of Goteborg]. Goteborgs Naturhistoriska Museum Arstryck 1984:31–45.
Hansen, J. M. 1977. Sedimentary history of the island Læsø, Denmark. Bulletin of theGeological Society of Denmark 26 (3–4):217–236.
Houmark-Nielsen, M., and K. H. Kjær. 2003. Southwest Scandinavia, 40–15 kyr BP:Palaeogeography and environmental change. Journal of Quaternary Science 18:769–786.
Houmark-Nielsen, M., J. Kruger and K. H. Kjær. 2005. De seneste 150.000 ar i Danmark.Istidslandskabet og naturens udvikling [The last 150,000 years in Denmark. The iceage landscape and the evolution of nature]. Geoviden–Geologi og Geografi 2:1–19.
Kinze, C. C. 1995. Danish whale records 1575–1991 (Mammalia, Cetacea). Review of whalespecimens stranded, directly or incidentally caught along the Danish coasts. Steenstrupia21:155–196.
Kinze, C. C., S. Tougaard and H. J. Baagøe. 1998. Danske hvalfund i perioden 1992–1997[Danish whale records 1992–1997]. Flora og Fauna 104:41–53.
Kinze, C. C. 2006a. Hvaler i Kolding Fjord [Whales in Kolding Fiord]. Koldingbogen2006:171–181.
Kinze, C. C. 2006b. Ny hval for Danmark: Brydeshval (Balaenoptera brydei) stranded vedKyndby i Isefjord [A new whale in Denmark: Bryde’s whale (Balaenoptera brydei) strandedat Kyndby in Isefiord]. Flora og Fauna 112:61–66.
Kinze, C. C. 2007. Hvaler [Whales]. Pages 264–311 in H. J. Baagøe and T. S. Jensen, eds.Dansk Pattedyr Atlas [Atlas of Danish mammals]. Gyldendal, Copenhagen, Denmark.
280 MARINE MAMMAL SCIENCE, VOL. 26, NO. 2, 2010
Kinze, C.C., H. J. Baagøe and S. Tougaard. 2001. Fund af eksotiske hvaler og delfiner IDanmark: Hyppighed og herkomst [Finds of exotic whales and dolphins in Denmark:Frequency and area of origin]. Flora og Fauna 107:107–116.
Lepiksaar, J. 1966. Zahnwalfunde in Schweden. Bijdragen tot de Dierkunde 36:3–16.Lepiksaar, J. 1986. The Holocene History of Theriofauna in Fennoscandia and Baltic Coun-
tries. Striae 24:51–70.Liljeborg, W. 1861. Ofversigt af de inom Skandinavien (Sverige och Norrige) antraffade
Hvalartade Daggdjur (Cetacea) [Review of whale species (Cetacea) occurring in Scandi-navia (Sweden and Norway)]. Upsala Universitets Arsskrift. 118 pp.
Liljeborg, W. 1867. On two subfossil whales discovered in Sweden. Nova Acta of the RoyalSociety of Sciences at Upsala, Series III, Volume VI. 48 pp.
Liljeborg, W. 1874. Sveriges och Norges Daggdjur [The mammals of Sweden and Norway].Upsala, Sweden.
Liljegren, R. 1975. Subfossila vertebratfynd fran Skane [Subfossil vertebrate remains fromSkane]. Report 8, Department of Quaternary Geology, University of Lund, Lund,Sweden. 187 pp.
Liljegren, R., and P. Lageras. 1993. Fran mammutstapp till kohage. Djurens historia i Sverige[From mammoth steppe to cultural steppe. The history of animals in Sweden]. Wallin& Dalholm Boktryckeri AB, Lund, Sweden.
Møhl, U. 1970. Fangstdyrene ved de danske strande [Seal and whale hunting on the Danishcoasts]. KUML. Arbog for Jysk arkæologisk Selskab 1970:297–329.
Nilsson, S. 1847. Skandinaviens Fauna I. Daggdjuren [The fauna of Scandinavia I. Themammals]. C. W. K. Gleerups Forlag, 2nd edition. Lund, Sweden.
Nybelin, O. 1942. Gammalt och nytt om Swedenborgsvalen [About the Swedenborg whale].Meddelanden fran Goteborgs Musei Zoologiska Avdeling 101:13–28.
Nybelin, O. 1946. Gronlandsvalen fran Guldheden och Swedenborgsvalen fran Landerigatan[The Greenland whale from Guldheden and the Swedenborg whale from Landerigatan].Goteborgs Museii Arstryck:103–115.
Petersen, K. S. 2004. Late Quaternary environmental changes recorded in the Danish ma-rine molluscan faunas. Geological Survey of Denmark and Greenland Bulletin 3:1–268.
Petersen, K. S., and A. Buch. 1974. Dislocated tills with Paleogene and Pleistocene marinebeds. Tectonics, lithology, macro- and microfossils. Danmarks geologiske Undersøgelser,arbog 1973:63–91.
Rasmussen, K. L. 2000. 14C-dateringer, København 1999 [Radiocarbon datings, Copen-hagen 1999]. Arkæologiske udgravninger i Danmark 1999, Det Arkæologiske Nævn,København 314–325.
Rasmussen, K. L., J. L. Boldsen, H. K. Kristensen, L. Skytte, K. L. Hansen, L. Mølholm,P. Grootes and M.-J. Nadeau. 2008. Mercury levels in Danish Medieval human bones.Journal of Archaeological Science 35:2295–2306.
Rasmussen, K. L., P. Bjerregaard, P. H. Gommesen and O. L. Jensen. 2009. Arsenic in Danishand Swedish Mesolithic and Neolithic human bones—diet or diagenesis? Journal ofArchaeological Science 36:2826–2834.
Reimer, P. J., M. G. L. Baillie, E. Bard, A. Bayliss, J. W. Beck, C. Bertrand, P. G. Blackwell,C. E. Buck, G. Burr, K. B. Cutler, P. E. Damon, R. L. Edwards, R. G. Fairbanks, M.Friedrich, T. P. Guilderson, K. A. Hughen, B. Kromer, F. G. McCormac, S. Manning,C. Bronk Ramsey, R. W. Reimer, S. Remmele, J. R. Southon, M. Stuiver, S. Talamo,F. W. Taylor, J. Van Der Plicht and C. E. Weyhenmeyer. 2004. IntCal04 terrestrialradiocarbon age calibration, 0–26 cal kyr BP. Radiocarbon 46:1029–1058.
Sommer, R. S., J. Pasold and U. Schmolcke. 2008. Post-Glacial immigration of the harbourporpoise (Phocoena phocoena) into the Baltic Sea. Boreas 37:458–464.
Stenstrop, G. 1994. En hval fra stenalderhavet [A whale from the stone age sea]. GeologiskNyt 3:6–8.
Tauber, P. 1878–1892. Zoologica Danica. 1. Bind. Pattedyr. Copenhagen, Denmark.
AARIS-SØRENSEN ET AL.: SUBFOSSIL WHALE BONES 281
Trolle-Lassen, T. 1985. En zooarkæologisk analyse af Ertebøllepladsen Tybrind Vig, primærtbaseret pa knogler af pelsdyr og kronhjort [A zooarchaeological analysis of the bonematerial from the Ertebølle site of Tybrind Vig]. M.Sc. thesis, University of Aarhus,Aarhus, Denmark. 168 pp.
Trueman, C. N., A. K. Behrensmeyer, N. Tuross and S. Weiner. 2004. Mineralogical and com-positional changes in bones exposed on soil surfaces in Amboseli National Park, Kenya:Diagenetic mechanisms and the role of sediment pore fluids. Journal of ArchaeologicalScience 31:721–739.
Winge, H. 1899. Om nogle Pattedyr i Danmark [About some mammals in Denmark].Videnskabelige Meddelelser fra Dansk naturhistorisk Forening 51:283–316.
Winge, H. 1904. Om jordfundne Pattedyr fra Danmark [About subfossil remains of mammalsfrom Denmark]. Meddelelser fra Dansk naturhistorisk Forening 56:193–304.
Received: 28 February 2009Accepted: 25 July 2009