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

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https://doi.org/10.1111/j.1748-7692.2009.00356.x

https://doi.org/10.1111/j.1748-7692.2009.00356.x

https://doi.org/10.1111/j.1748-7692.2009.00356.x

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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


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


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,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


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-


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


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


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


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


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.,


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


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


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,


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.


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.


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


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


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


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

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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


LuH

fTa

Th

Lab

no.

�g/

g�

g/g

�g/

g�

g/g

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700

0.11

41.

860

0.10

900.

880

KLR

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10.

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0.05

6nd

0.10

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LR-1

702

0.14

70.

585

0.07

410.

755

KLR

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30.

074

2.26

00.

1690

0.96

9K

LR-1

704

0.42

00.

229

nd1.

610

KLR

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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

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90.

028

1.40

00.

1140

0.97

1K

LR-1

710

nd0.

318

nd0.

047

KLR

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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

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714

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0.22

30.

0253

0.22

6


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,


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.


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


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.

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Received: 28 February 2009Accepted: 25 July 2009

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Late Pleistocene and Holocene whale remains (Cetacea) from