Deciduous Tusks and Small Permanent Tusks of the Woolly Mammoth, Mammuthus primigenius (Blumenbach, 1799) Found on Beaches in The Netherlands - MDPI

 
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Deciduous Tusks and Small Permanent Tusks of the Woolly Mammoth, Mammuthus primigenius (Blumenbach, 1799) Found on Beaches in The Netherlands - MDPI
Article
Deciduous Tusks and Small Permanent Tusks of the
Woolly Mammoth, Mammuthus primigenius
(Blumenbach, 1799) Found on Beaches in
The Netherlands
Dick Mol 1, *, André Bijkerk 2 and Joanne P. Ballard 3
 1   Natural History Museum Rotterdam, 3015 AA Rotterdam, The Netherlands
 2   Independent Researcher, 2726 CP Zoetermeer, The Netherlands; andrebijkerk@gmail.com
 3   Tennessee Valley Authority, Knoxville, TN 37902, USA; jpballard0@tva.gov
 *   Correspondence: dickmol@telfort.nl

 Academic Editor: Maria Rita Palombo
                                                                                                   
 Received: 25 March 2018; Accepted: 22 May 2018; Published: 28 May 2018                            

 Abstract: Tusks of very young mammoths and other proboscideans are extremely rare in the fossil
 record. This article presents all deciduous tusks and newly-developing permanent tusks of woolly
 mammoths which are known from Dutch localities and the North Sea. Four deciduous tusks are from
 The Netherlands and one is from Germany. Five specimens of newly-developing permanent tusks
 are also discussed. In this article, we present several characteristics and details of those tusks such as
 presence/absence of roots, and cementum and enamel layers that should facilitate an identification
 when new findings are made. We also describe a dental element, a digitelle, that is easily mistaken for
 a tusk of a juvenile woolly mammoth.

 Keywords: proboscideans; dentition; milk tusk and permanent tusks; digitelle; North Sea; Maasvlakte;
 Zandmotor; Rockanje

1. Introduction
      The North Sea floor, located between the British Isles and the continent of Europe, is an important
source of fossil mammal remains from the entire Quaternary [1]. Several mammalian faunas inhabited
the area during various episodes of the Pleistocene. The terrestrial mammals from the North Sea
are divided into four fauna-associations which are regarded as groups of larger mammals with a
comparable age, although not necessarily regarded as contemporaneous [2].
      The oldest, Early to Middle Pleistocene Fauna-associations I and II, contain heavily mineralized
and very dark fossils including the proboscideans Anancus arvernensis (Croizet and Jobert, 1828),
Mammuthus meridionalis (Nesti, 1825) and Mammuthus trogontherii (Pohlig, 1885). Fauna-association
III is much younger, of Late Pleistocene age, with Elephas antiquus (Falconer and Cautley, 1847)
and Mammuthus primigenius. The bulk of the radiocarbon dates fall within Maritime Isotope
Stage 3 (57–29 Ka cal BP). Fauna-association IV is Holocene, and does not contain any remains
of proboscideans.
      The fossils of these mammals have routinely been brought up in the nets of fishing vessels such
as beam trawlers [3,4]. One of the best known and richest sources of fossils in the Southern Bight of
the North Sea is the Eurogeul area off the coast of the province of South Holland. These fossils are
found in fluvial sediments deposited in the former deltas of the Rhine and Meuse Rivers, before they
were flooded by rising sea levels at the end of the last ice age [5,6]. The Eurogeul, or Eurogully, is a
channel for large ships to enter the ports of Rotterdam (the Euro-Meuse channel). This channel fills

Quaternary 2018, 1, 7; doi:10.3390/quat1010007                             www.mdpi.com/journal/quaternary
Deciduous Tusks and Small Permanent Tusks of the Woolly Mammoth, Mammuthus primigenius (Blumenbach, 1799) Found on Beaches in The Netherlands - MDPI
Quaternary 2018, 1, 7                                                                                              2 of 14

Quaternary 2018, 2, x FOR PEER REVIEW                                                                             2 of 14
in with sediment, which must routinely be removed to maintain an adequate channel depth for the
passage  of ships.
passage of  ships. The
                     The fossil-rich
                          fossil-rich sediments
                                       sediments are
                                                   are dredged
                                                       dredged and
                                                                 and deposited
                                                                      deposited along
                                                                                  along the
                                                                                          the coastline
                                                                                               coastline ofof The
                                                                                                              The
Netherlands, which
Netherlands,  which reinforces
                      reinforces and
                                   and extends
                                        extends the
                                                 the beaches.
                                                     beaches. The
                                                              The same
                                                                  same process
                                                                        process isis used
                                                                                     used for
                                                                                           for the
                                                                                                theconstruction
                                                                                                    construction
of Maasvlakte   2, an artificial extension  of the Rotterdam   harbor (Figure 1).  This results
of Maasvlakte 2, an artificial extension of the Rotterdam harbor (Figure 1). This results in the  in the creation
                                                                                                         creation
of new localities  where  the  remains   of Pleistocene and Holocene   animals  may    be
of new localities where the remains of Pleistocene and Holocene animals may be discovered. discovered.

     Figure 1. Map of the Eurogeul area that ships pass through on their approach to Rotterdam. Pale
      Figure 1. Map of the Eurogeul area that ships pass through on their approach to Rotterdam. Pale yellow
     yellow represents the dredged area. The popular Maasvlakte 2 fossil-hunting beach is marked in
      represents the dredged area. The popular Maasvlakte 2 fossil-hunting beach is marked in medium
     medium yellow—this is where the dredged sediment is deposited on the beach. The green silhouette
     yellow—this is where the dredged sediment is deposited on the beach. The green silhouette in the
     in the upper right is The Netherlands with the Eurogeul area marked with a red dot.
      upper right is The Netherlands with the Eurogeul area marked with a red dot.

      In recent years, many private collectors have created collections of various fossils found on the
      In recent
seashore.   Theseyears,
                   beaches many
                              are aprivate   collectors
                                    rich source            have created
                                                   of well-preserved       collections
                                                                         skeletal        oflarge
                                                                                  parts of   various   fossils mammals
                                                                                                  terrestrial  found on
the  seashore.    These    beaches   are  a rich   source    of well-preserved     skeletal
such as woolly mammoths and woolly rhinoceroses of the mammoth fauna. However, remains of    parts  of  large  terrestrial
mammals
small         such as
       terrestrial,     woolly
                    marine,    andmammoths
                                    freshwaterand      woolly
                                                   aquatic       rhinoceroses
                                                             mammals     such asofbeavers
                                                                                    the mammoth
                                                                                            and ottersfauna.
                                                                                                         have However,
                                                                                                               also been
remains    of small  terrestrial,  marine,  and   freshwater    aquatic   mammals    such   as beavers
collected. Other interesting finds include insect remains, birds, fish, and even coprolites of hyenas    and  otters have
                                                                                                                      [7–
also  been   collected.  Other   interesting   finds  include    insect remains,  birds,  fish, and
14]. These fossil finds are often well documented, and reveal many previously unknown details about  even   coprolites  of
hyenas    [7–14].  These   fossil finds are  often   well   documented,     and  reveal  many    previously
the individuals and their species, as well as their habitat and faunal assemblages. Through amateur            unknown
detailscollecting,
beach    about theimportant
                       individuals    and have
                                  insights  their been
                                                    species,
                                                          made asfor
                                                                   well
                                                                     theas  their habitat
                                                                          deciduous   and theandvery
                                                                                                  faunal
                                                                                                      smallassemblages.
                                                                                                             permanent
Through     amateur    beach  collecting, important      insights  have  been
tusks of woolly mammoths. Little was known about these deciduous tusks and the made   for the  deciduous    and  theearly
                                                                                                                     very
small   permanent     tusks  of woolly   mammoths.
development of the permanent tusks of these proboscideans.Little was  known    about  these   deciduous    tusks  and the
earlyThe
       development      of the  permanent     tusks   of these  proboscideans.
           juvenile deciduous and permanent tusks described here are identified as belonging to the
      The   juvenile deciduous
late Pleistocene-age     Mammuthus  andprimigenius,
                                         permanent based tusks described    here are identified
                                                                 on their morphology       [15–17].asEarly
                                                                                                       belonging   to the
                                                                                                            and middle
late Pleistocene-age     Mammuthus       primigenius,     based   on their morphology      [15–17].
Pleistocene faunal remains from the bottom of the North Sea can easily be distinguished from late    Early   and  middle
Pleistocene remains
Pleistocene    faunal remains
                         by theirfrom    themineralized
                                   heavily    bottom of the       North
                                                              state      Sea can easily
                                                                    of preservation       be distinguished from late
                                                                                      [1,8,9,18].
Pleistocene remains by their heavily mineralized state of preservation [1,8,9,18].
2. Literature Review for Deciduous and Permanent Tusks
     Several publications describe details about deciduous and permanent tusks of Mammuthus
primigenius [10,15–17,19,20]. However, the number of reported specimens is limited.
Deciduous Tusks and Small Permanent Tusks of the Woolly Mammoth, Mammuthus primigenius (Blumenbach, 1799) Found on Beaches in The Netherlands - MDPI
Quaternary 2018, 1, 7                                                                                                      3 of 14

2. Literature Review for Deciduous and Permanent Tusks
     Several publications describe details about deciduous and permanent tusks of Mammuthus
primigenius [10,15–17,19,20]. However, the number of reported specimens is limited.
     Rountrey et al. [17] stressed the importance of finding additional tusk specimens in their landmark
research on early tooth development, gestation, and season of birth in mammoths regarding these
dental elements. In their conclusion they stated:

      This work shows the potential value of detailed individual life history studies using
      teeth. With work on additional specimens, these techniques could lead to construction
      of an accurate tooth development schedule for mammoths, and this information is critical
      for studies of mammoth life history that will help elucidate ecological factors associated
      with extinction.

     Every find of deciduous tusks and small permanent tusks of extremely young mammoths can
contribute greatly to our scientific understanding in this area of study, and should be recorded.
Hence, the amateur collectors of fossil remains on Dutch beaches deserve support and should be
encouraged both to learn to recognize these rare specimens and to have opportunities to present them
to paleontologists for examination. This paper intends to contribute to this effort, with a short overview
of the literature and a discussion of both the known deciduous tusks and the smallest permanent
tusks of juvenile mammoths from the environment of The Netherlands, which have been identified in
recent years.
     There are other dental elements and parts thereof that can be easily confused with the smallest
tusks. One of these confounding fossils is presented here, a fossil digitelle (“finger”) of a plate of a
premolar of a juvenile woolly mammoth, which was considered to be a possible small tusk. Mammoths
and modern elephants go through a series of six sets of adult molars in their lifetime. The molars are
classified with capital letters for upper molars and lower case letters for lower jaw molars (Table 1).

      Table 1. The dental formulae (following Sikes [21]) used for Mammuthus primigenius and recent African
      and Asian elephants for each side.

                                          1I                 0C                       3 dP                       3M
       Maxilla (Upper Jaw)
                                     (1× Incisive)       (No Canines)        (3 Deciduous Premolars)          (3 Molars)
                                         0i                    0c                      3 dp                      3m
     Mandibula (Lower jaw)
                                    (No incisives)        (no canines)       (3 deciduous premolars)          (3 molars)
      The uppercase letters I, C, P, and M are used to designate the upper jaw; and the lowercase letters i, c, p, and m are
      used for the lower jaw.

     Recognizing and determining fossils to species level requires comparison with known specimens.
Because of the rarity and the difficulty in recognizing the deciduous and juvenile permanent tusks
of Mammuthus primigenius, the fossil record is very limited. Sikes [21] discussed details about these
teeth from the African elephant, Loxodonta africana (Blumenbach, 1797), providing useful specimens
for comparison. Mashchenko [15] described several of these juvenile tusks, and concluded that both
deciduous tusks and permanent tusks develop in the fetus during the earliest stages simultaneously
and that the former were merely rudimentary without any function, being quickly replaced by the
permanent tusks.
     Rountrey et al. [17] published a detailed analysis of two extraordinary late Pleistocene juvenile
mammoth mummies from Siberia, giving a comprehensive description of their dental elements.
One complete mummy was Lyuba from Yamal, a female individual who died when she was one month
old. Remarkably, she was missing both permanent tusks. The other was Oimyakon from Yakutia,
a partial mummy with the right permanent tusk, the left one being lost. Its gender is unknown and
it was approximately seven months old at time of death. It had already lost the deciduous tusks,
described and photographed by Maschenko et al. [16]. In cross-section, the oval-shaped permanent
Deciduous Tusks and Small Permanent Tusks of the Woolly Mammoth, Mammuthus primigenius (Blumenbach, 1799) Found on Beaches in The Netherlands - MDPI
Quaternary 2018, 1, 7                                                                                              4 of 14

tusk measured 76 mm in length and was entirely covered with cementum and characterized by the tip
of the tusk being covered with an enamel cap.
      Van der Plicht et al. [10] described a somewhat larger juvenile skull with a right permanent tusk
Quaternary 2018, 2,in
still anchored        the alveolus.
                    x FOR           This skull was trawled from the North Sea floor in the Eurogeul.4 of
                          PEER REVIEW                                                                 It 14
                                                                                                         is
mentioned here to illustrate the typical black enamel, which eventually becomes worn down due to
usage (Figure 2). TheseThese publications
                              publications are
                                           are useful
                                               useful references
                                                      references for identifying
                                                                     identifying and analyzing
                                                                                     analyzing these rare
dental elements.

      Figure
      Figure 2.
             2. Incomplete
                Incomplete cranium
                            cranium of of aa juvenile
                                             juvenile Mammuthus
                                                      Mammuthus primigenius
                                                                 primigenius with
                                                                             with the
                                                                                   the right
                                                                                        right permanent
                                                                                              permanent tusk
                                                                                                           tusk in
                                                                                                                in
      anatomical  position. For  details   and  measurements   see Van  der Plicht  et  al. [10]. Locality: Euro-
      anatomical position. For details and measurements see Van der Plicht et al. [10]. Locality: Euro-Maasgeul,
      Maasgeul,
      North Sea. North  Sea. Collection:
                  Collection: Klaas Post,Klaas   Post, Urk.
                                            Urk. Photo: HansPhoto: Hans Wildschut.
                                                              Wildschut.

3. Review of Deciduous and Permanent Tusk Collections in The Netherlands
3. Review of Deciduous and Permanent Tusk Collections in The Netherlands
     The
     The following
         following is
                    is aa brief
                           brief overview
                                 overview of   all known
                                            of all known fossil
                                                         fossil deciduous
                                                                deciduous and permanent tusks
                                                                          and permanent tusks of
                                                                                              of
Mammuthus   primigenius  in or close to The Netherlands.
Mammuthus primigenius in or close to The Netherlands.

3.1.
3.1. Terra Maris Museum
     Terra Maris Museum Collection
                        Collection in
                                   in Oostkapelle
                                      Oostkapelle
      A
      A deciduous
        deciduous tusk
                     tusk in
                           in the
                              the collection
                                  collection of
                                             of the
                                                the Museum
                                                    Museum Terra
                                                               Terra Maris
                                                                     Maris (Oostkapelle)
                                                                            (Oostkapelle) was
                                                                                           was found
                                                                                                found on on aa trail
                                                                                                               trail
consisting
consisting ofof sea
                sea shells
                    shells in
                            in Zeeland
                               Zeeland near
                                         near Veere  in the
                                              Veere in  the late
                                                            late 1980s.
                                                                  1980s. The
                                                                         The very
                                                                             very small
                                                                                   small and
                                                                                         and fragile
                                                                                              fragile specimen
                                                                                                        specimen
shows
shows the
        the typical
            typical well-developed
                     well-developed root
                                       root and
                                            and has
                                                 has been
                                                     been described
                                                           described by
                                                                      by Langeveld   and Mol
                                                                          Langeveld and   Mol [22].
                                                                                               [22]. It
                                                                                                     It belonged
                                                                                                        belonged
to
to a very young Mammuthus primigenius, probably only a couple of months old at the time ofits
   a very young    Mammuthus     primigenius, probably   only a  couple  of months old at the time  of   itsdeath.
                                                                                                             death.

3.2. Collection of Jan van der Steeg
     A perfectly
        perfectlypreserved
                    preserved    deciduous
                              deciduous    tusktusk
                                                 was was
                                                      foundfound
                                                              in the in  the Holt-und-Haar
                                                                     Holt-und-Haar       sand pit,sand
                                                                                                    close pit,   close to
                                                                                                           to Gildehaus
Gildehaus
in Germanyinnear
               Germany     nearborder.
                   the Dutch     the Dutch
                                         It isborder.  It is catalogued
                                               catalogued                  as number
                                                             as number P00170       in theP00170   in theofcollection
                                                                                            collection       Jan van derof
Jan van
Steeg inder  SteegMeijer
         Losser.   in Losser.   Meijer [23]more
                          [23] provided      provided
                                                  detailsmore  details
                                                           about        about
                                                                  the site  andthe
                                                                                 its site and its mammalian
                                                                                     mammalian       fauna. Thefauna.
                                                                                                                    entire
The  entire is
specimen    specimen
               somewhatis somewhat
                            club shapedclubhasshaped
                                                 a veryhasweak
                                                            a veryS-curve.
                                                                   weak S-curve.      The specimen
                                                                              The specimen              was described
                                                                                                  was described       and
and  photographed
photographed          by Mol
                 by Mol         and Boeskorov
                          and Boeskorov      [24]. [24]. No traces
                                                   No traces         of wear
                                                               of wear         are visible,
                                                                         are visible,         indicating
                                                                                        indicating   that itthat
                                                                                                              mustit must
                                                                                                                     have
have  been embedded
been embedded     entirelyentirely  in the alveolus.
                            in the alveolus.   Mol andMol     and Boeskorov
                                                         Boeskorov                [24] suggested
                                                                       [24] suggested                 that this
                                                                                          that this animal    mustanimal
                                                                                                                     have
must  have died
died before       before
              the age     the year.
                      of one   age ofThe
                                       onetusk
                                            year.   The tusk69.9
                                                 measures      measures
                                                                  mm in 69.9    mmThe
                                                                           length.     in length.   The tipwith
                                                                                           tip is covered     is covered
                                                                                                                    a thin
with a thin enamel layer which in turn is covered by a very thin layer of cementum. The diameter of
the enamel cap is between 7.1 and 14 mm.

3.3. Collection of Walter Langendoen
   Another right deciduous tusk of a Mammuthus primigenius was collected on May 2014 on the
man-made beach of Maasvlakte 2 and added as specimen number WL0181 to the collection of Walter
Deciduous Tusks and Small Permanent Tusks of the Woolly Mammoth, Mammuthus primigenius (Blumenbach, 1799) Found on Beaches in The Netherlands - MDPI
Quaternary 2018, 1, 7                                                                                                    5 of 14

enamel layer which in turn is covered by a very thin layer of cementum. The diameter of the enamel
cap is between 7.1 and 14 mm.
         Quaternary 2018, 2, x FOR PEER REVIEW                                                                        5 of 14
3.3. Collection of Walter Langendoen
        reasonably
     Another    rightwell preserved.tusk
                       deciduous      Its tipofconsists of dentin primigenius
                                                a Mammuthus        covered by a was
                                                                                  cap ofcollected
                                                                                          enamel. The
                                                                                                    on root
                                                                                                        Mayis2014
                                                                                                               damaged
                                                                                                                     on the
        in the area of the pulp cavity and the exact original length cannot be determined. This specimen is
man-made beach of Maasvlakte 2 and added as specimen number WL0181 to the collection of Walter
        practically straight and has a preserved length of 46 mm. The enamel cap’s length is 17 mm and its
Langendoen (Figure 3). It is currently on display in the Historyland museum in Hellevoetsluis.
        diameter varies between 6 and 9 mm.
Mol et al. [9], Langeveld
              The             and
                  original tusk wasMol   [22]
                                      used   to and
                                                makeMolsome [18] described
                                                              casts.          this small
                                                                     Unfortunately,        deciduous
                                                                                      it broke           tusk,the
                                                                                                in half along   which
                                                                                                                  lengthwas
reasonably
        of thewell
               axispreserved.   Its tip consists
                    during the production     of theof  dentin
                                                     casting     covered
                                                              mold.        by a cap
                                                                      However,        of enamel.
                                                                                 this accident      The root
                                                                                                 provided      is damaged
                                                                                                           insight in the
in the area  of the
        interior,    pulp
                  which     cavity
                          shows  theand   the exact
                                     different   toothoriginal    lengthindicating
                                                         tissues clearly   cannot bedentin,
                                                                                        determined.
                                                                                               cementum  This specimen
                                                                                                            and  enamel. is
practically  straight
        The depth      andstill-open
                    of the   has a preserved
                                      pulp cavity,length    of 46 mm.
                                                      the volume         The enamel
                                                                     of dentin and the cap’s    length
                                                                                         cementum    andisenamel
                                                                                                            17 mmcover
                                                                                                                    and its
        are visible in Figure 3H.
diameter varies between 6 and 9 mm.The  tusk   was eventually    restored.

              Figure 3. Right deciduous tusk of Mammuthus primigenius. Locality: Beach of Maasvlakte. Collection:
      Figure 3. Right deciduous tusk of Mammuthus primigenius. Locality: Beach of Maasvlakte. Collection:
              Walter Langendoen (Hellevoetsluis) WL0181. Scale bar: 5 cm. Photo: Hans Wildschut. (A) Medial
      Walter Langendoen (Hellevoetsluis) WL0181. Scale bar: 5 cm. Photo: Hans Wildschut. (A) Medial view;
              view; (B) cranial view; (C) lateral view; (D) anterior view conical cap (tip of the tusk); (E) posterior
      (B) cranial view; (C) lateral view; (D) anterior view conical cap (tip of the tusk); (E) posterior view,
              view, circumference of the root including the cross section of the pulp cavity; (F) interior view after
      circumference
              breakageof  the root
                        medial        including
                               half; (G)           the after
                                         interior view  crossbreakage
                                                              sectionlateral
                                                                       of thehalf;
                                                                               pulpandcavity;  (F)ofinterior
                                                                                       (H) details            view
                                                                                                     the anterior    after
                                                                                                                  view.
      breakage    medial half;
              c: cementum;      (G) interior
                            d: dentine;        view
                                        e: enamel;  pc:after
                                                        pulpbreakage   lateral
                                                             cavity. Photo: Hanshalf; and (H) details of the anterior
                                                                                  Wildschut.
      view. c: cementum; d: dentine; e: enamel; pc: pulp cavity. Photo: Hans Wildschut.
Deciduous Tusks and Small Permanent Tusks of the Woolly Mammoth, Mammuthus primigenius (Blumenbach, 1799) Found on Beaches in The Netherlands - MDPI
Quaternary 2018, 1, 7                                                                                                       6 of 14

     The original tusk was used to make some casts. Unfortunately, it broke in half along the length
of the axis during the production of the casting mold. However, this accident provided insight in
the interior, which shows the different tooth tissues clearly indicating dentin, cementum and enamel.
The depth of the still-open pulp cavity, the volume of dentin and the cementum and enamel cover are
 Quaternary 2018, 2, x FOR PEER REVIEW                                                           6 of 14
visible in Figure 3H. The tusk was eventually restored.

 3.4.Collection
3.4.  CollectionofofPeter
                     PeterSoeters
                           Soeters

      AAcomplete
           completeright
                       rightdeciduous
                              deciduoustusktuskwas
                                                 wasfound
                                                      foundon onthe
                                                                  thebeach
                                                                      beachof ofRockanje
                                                                                 Rockanje(South
                                                                                            (SouthHolland
                                                                                                     HollandProvince)
                                                                                                                 Province)
ininMay
     May20082008and
                 andititwas
                         wasadded
                               addedtotothe
                                          thecollection
                                              collectionofofPeter
                                                             PeterSoeters
                                                                    SoetersininDelft
                                                                                Delft(Figure
                                                                                      (Figure4).
                                                                                               4).Its
                                                                                                   Itsroot
                                                                                                       rootisissomewhat
                                                                                                                somewhat
  curved    and  has  a  rather  small   pulp  cavity. The   tusk  is slightly  incurvate   as well
curved and has a rather small pulp cavity. The tusk is slightly incurvate as well and has a pristine  and  has   a pristine
                                                                                                                      cap
ofcap  of black
    black         enamel.
            enamel.  It has Ita shallow,
                                has a shallow,   weakly-developed
                                          weakly-developed       groovegroove     on the medial
                                                                         on the medial    side. Theside. The maximum
                                                                                                      maximum      length
islength
    52 mm  is and
              52 mmthe and  the thickness
                       thickness    is 6 mmismeasured
                                                6 mm measured       in the middle
                                                           in the middle             of the flattened
                                                                           of the flattened    root. Theroot.  The crown
                                                                                                           crown    has a
  has  a lateral length   of  9 mm   and   a medial
lateral length of 9 mm and a medial length of 7 mm.  length   of 7 mm.

      Figure 4. Right deciduous tusk of Mammuthus primigenius. Locality: Beach of Rockanje. Collection:
      Peter Soeters
       Figure        (Delft).
              4. Right        Scale bar:
                        deciduous   tusk 5ofcm. Photo: Hans
                                             Mammuthus       Wildschut:
                                                        primigenius.     (A) Medial
                                                                     Locality:       view;
                                                                               Beach of    (B) lateral
                                                                                        Rockanje.      view;
                                                                                                  Collection:
      and (C)Soeters
       Peter  cranial(Delft).
                      view. Scale bar: 5 cm. Photo: Hans Wildschut: (A) Medial view; (B) lateral view; and
       (C) cranial view.
3.5. Collection of Donny Chrispijn
 3.5. Collection of Donny Chrispijn
      A complete right deciduous tusk was found by Peter Soeters on the beach of Rockanje in April
       A complete
2008, and    it was addedrighttodeciduous      tuskofwas
                                  the collection            found
                                                        Donny       by Peter
                                                                 Chrispijn   inSoeters
                                                                                Hoek van onHolland
                                                                                             the beach      of Rockanje
                                                                                                         (Figure  5). The in    April
                                                                                                                            typical
 2008, and
groove          it wason
          is visible      added    to theside,
                             the medial     collection    of Donny
                                                  facilitating         Chrispijn in Hoek
                                                                 the determination.             van Holland
                                                                                          The smaller      length(Figure
                                                                                                                    compared 5). The
                                                                                                                                  to
 typical
the         groove
     specimen          is visible
                   in Section     3.4on    the medial
                                      is noted,    whichside,     facilitating
                                                            is probably    due totheresorption
                                                                                      determination.         The smaller
                                                                                                  in the growing       processlength
                                                                                                                                  of
 compared
the permanent   to the
                    tuskspecimen
                             behind init. Section   3.4 is length
                                          The original     noted, must
                                                                   whichhaveis probably    due to resorption
                                                                                 been distinctly     longer. This in the
                                                                                                                      wouldgrowing
                                                                                                                                also
 process
mean    thatofthe
                thepermanent
                     permanenttusk   tuskwasbehind    it. The
                                                 present       original
                                                            during        length
                                                                     the first    must
                                                                               year       have
                                                                                      of the     been
                                                                                              calf’s  lifedistinctly  longer.
                                                                                                            [16]. Sikes          This
                                                                                                                          [21] also
 would     also  mean     that  the permanent       tusk  was   present  during    the first year
described this growing process for African elephants. The maximum length of this specimen is 37 mm of   the  calf’s life [16].  Sikes
 [21]the
and    also  described
          thickness     is 7this
                              mmgrowing
                                   measured  process
                                                 in thefor  African
                                                         middle       elephants.
                                                                  of the flattenedThe    maximum
                                                                                     root.  The crown   length   of this the
                                                                                                           has almost     specimen
                                                                                                                               same
 is 37 mm and
dimensions           the
                 as in      thickness
                        Section          is 7 amm
                                   3.4, has          measured
                                                lateral  length ofin 9the
                                                                       mm  middle
                                                                             and a of   the flattened
                                                                                     medial    length ofroot.7 mm.TheThecrown
                                                                                                                           enamel has
 almost
cap        thepreserved
     is well     same dimensions          as in is
                              and its width      Section
                                                   12 mm3.4,    has a lateral
                                                             laterally  and 11length     of 9 mm and a medial length of 7
                                                                                mm medially.
 mm. The enamel cap is well preserved and its width is 12 mm laterally and 11 mm medially.
Deciduous Tusks and Small Permanent Tusks of the Woolly Mammoth, Mammuthus primigenius (Blumenbach, 1799) Found on Beaches in The Netherlands - MDPI
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         Figure
      Figure  5. 5.  Right
                  Right     deciduoustusk
                         deciduous     tuskofofMammuthus
                                                Mammuthus primigenius.
                                                              primigenius. Locality:
                                                                           Locality:Beach
                                                                                     BeachofofRockanje.
                                                                                               Rockanje.Collection:
                                                                                                          Collection:
         Donny
        Figure
      Donny      5.Chrispijn
               Chrispijn     (Hoekvan
                    Right deciduous
                           (Hoek    van  Holland).
                                      tusk          Scale
                                                     Scalebar:
                                            of Mammuthus
                                         Holland).             5 cm. Photo:
                                                            primigenius.
                                                           bar:  5 cm.      Hans
                                                                          Locality:
                                                                        Photo:     Wildschut.
                                                                                HansBeach     (A) Medial
                                                                                          of Rockanje.
                                                                                       Wildschut.         view; (B)
                                                                                                   (A) Collection:
                                                                                                        Medial  view;
         lateral
        Donny    view;  and
                 Chrispijn   (C)
                            (Hoekcranial
                                   van   view.
                                         Holland).
      (B) lateral view; and (C) cranial view.      Scale bar: 5 cm. Photo:  Hans  Wildschut. (A) Medial  view; (B)
        lateral view; and (C) cranial view.
   3.6. Collection of Hansjorg Ahrens
3.6. Collection of Hansjorg Ahrens
  3.6. Collection of Hansjorg Ahrens
         Hansjorg Ahrens from Leiden collected a small permanent right tusk of a very young woolly
      Hansjorg Ahrens from Leiden collected a small permanent right tusk of a very young woolly
   mammoth
        Hansjorg on Ahrens
                       the beach     of De
                                  from        Zandmotor,
                                           Leiden      collected which    is anpermanent
                                                                     a small      artificial extension
                                                                                               right tusk    ofof
                                                                                                                thea coastline
                                                                                                                      very young  at the  coast
                                                                                                                                       woolly
mammoth
   of Monster
  mammoth
              onon
                   the
                  (South beach
                            Holland
                     the beach
                                   of  DeProvince).
                                    of De
                                            Zandmotor,
                                            Zandmotor,
                                                                 which
                                                            Except
                                                                which    aissmall
                                                                     for is   an artificial   extension
                                                                                    indentation
                                                                             an artificial          on the
                                                                                            extension
                                                                                                              of  thecavity,
                                                                                                            of pulp
                                                                                                                       coastline
                                                                                                               the coastline    the  at the
                                                                                                                                     tusk
                                                                                                                                 at the    iscoast
                                                                                                                                         coast in
ofof
   Monster
   superb
     Monster   (South
            state          Holland
                    of preservation
                 (South                 Province).
                                           (Figure 6).Except
                           Holland Province).                Except
                                                             The tusk for
                                                                    for is a  small
                                                                           straight
                                                                        a small       indentation
                                                                                      and slightly
                                                                                   indentation            on  the
                                                                                                   on flattened     pulp    cavity,
                                                                                                                     giving an
                                                                                                        the pulp cavity,       theovalthe is in is
                                                                                                                                           tusk
                                                                                                                                    tuskcross-
insuperb
   superbstate
   section.  state
             It isof  of
                   coveredpreservation
                                in a thin
                       preservation           (Figure
                                             film
                                          (Figure            6). tusk
                                                    of6).black
                                                            The  The
                                                                 enameltusk
                                                                       is      is straight
                                                                            with
                                                                          straight the
                                                                                     and      and slightly
                                                                                        exception
                                                                                           slightly   of   a small
                                                                                                      flattened   flattened
                                                                                                                     sectionan
                                                                                                                    giving      giving
                                                                                                                                of        an oval
                                                                                                                                   the medial
                                                                                                                                  oval  cross-
cross-section.
   side which
  section.  It is isIttan
                        is colored.
                  covered  covered
                              in a thin  infilm
                                        The  alateral
                                                thin
                                                  of blackfilm   of blacksurface
                                                           and enamel
                                                                medial        enamel
                                                                           with          withtusk
                                                                                  theofexception
                                                                                         this   theofisexception
                                                                                                         arough
                                                                                                            smalldue   oftoaasmall
                                                                                                                    section    thin
                                                                                                                               of      section
                                                                                                                                       medial of
                                                                                                                                  theirregular
the  medial
   layerwhich
  side         side
          of enamel.    which
                 is tan The       is
                               typical
                          colored.    tan  colored.
                                       Thegroove
                                             lateralisand   The   lateral
                                                            visible  on the
                                                                medial      and   medial
                                                                              medial
                                                                         surface             surface
                                                                                        side.tusk
                                                                                    of this               of
                                                                                              Thisispermanent this  tusk
                                                                                                        rough duetusk       is rough
                                                                                                                        to a belonged    due
                                                                                                                                          to an
                                                                                                                              thin irregular    to a
thin irregular
   unborn
  layer  of enamel.layer
             or newborn  Theoftypical
                               enamel.
                              individual.    The
                                          grooveIt istypical
                                                      isthe      groove
                                                              smallest
                                                          visible           ismedial
                                                                         known
                                                                    on the     visible   on the
                                                                                   specimen
                                                                                       side.      medial
                                                                                                with
                                                                                             This              side.tusk
                                                                                                        a maximum
                                                                                                    permanent          This   permanent
                                                                                                                           length   of 37tomm.
                                                                                                                             belonged       an tusk
belonged
   The depth
  unborn     tonewborn
            or   an   unborn
                 of the   pulp     or  newborn
                                   cavity
                             individual.    is It
                                               22ismm. individual.
                                                      the               It is the
                                                              The maximum
                                                             smallest   known       smallestwith
                                                                                 circumference
                                                                                  specimen      knownata the specimen       with
                                                                                                               base of length
                                                                                                           maximum        the  pulpa maximum
                                                                                                                                   of cavity
                                                                                                                                      37 mm.is
length
  The   of 37
   43 mm;
       depth    mm.
             theof  the The
                         pulpdepth
                  diameter       is
                                  cavity ofisthe
                                    16 mm,    22andpulp
                                                  mm.         cavity
                                                        it measures
                                                             The      is922
                                                                  maximum  mm mm.fromThethemaximum
                                                                                circumferenceaxis to    thecircumference
                                                                                                    atthe    lateral
                                                                                                              base ofside.
                                                                                                                         the pulp at cavity
                                                                                                                                     the base is of
  43pulp
the  mm;cavity
            the diameter
                     is 43 mm; is 16   mm,
                                     the      and it measures
                                          diameter         is 16 mm,9and  mmitfrom     the axis
                                                                                 measures         to the
                                                                                               9 mm         lateral
                                                                                                         from    the side.
                                                                                                                      axis to the lateral side.

         Figure 6. Right permanent tusk of Mammuthus primigenius. Locality: Beach of de Zandmotor.
      Figure     Right
        Figure6. 6.
         Collection:    permanent
                       Hansjorg
                     Right       Ahrens
                            permanent      of Mammuthus
                                     tusktusk
                                          (Leiden).        primigenius.
                                                     Scale bar:
                                               of Mammuthus              Locality:
                                                                5 primigenius.
                                                                  cm. Photo:  Hans Beach of de Zandmotor.
                                                                                    Wildschut.
                                                                               Locality: Beach (A)
                                                                                                of Lateral  Collection:
                                                                                                           view;
                                                                                                   de Zandmotor. (B)
      Hansjorg
         medial  Ahrens
                 view;  (C)(Leiden).
                             cranial  Scale
                                     view;   bar:
                                            and   5
                                                (D) cm.  Photo:  Hans
                                                     circumference     Wildschut.
                                                                     pulp cavity.  (A) Lateral view; (B) medial
        Collection: Hansjorg Ahrens (Leiden). Scale bar: 5 cm. Photo: Hans Wildschut. (A) Lateral view; (B)      view;
      (C) cranial   view; and   (D) circumference    pulp  cavity.
        medial view; (C) cranial view; and (D) circumference pulp cavity.
Deciduous Tusks and Small Permanent Tusks of the Woolly Mammoth, Mammuthus primigenius (Blumenbach, 1799) Found on Beaches in The Netherlands - MDPI
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3.7.
3.7. Collection
     Collection of
                 of Dick
                    Dick Mol
                         Mol
      Peter
      Peter Soeters   from Delft
             Soeters from  Delft collected
                                 collectedaapermanent
                                              permanenttusk
                                                         tuskofofaajuvenile
                                                                     juvenilewoolly
                                                                              woollymammoth
                                                                                      mammoth  onon
                                                                                                  thethe beach
                                                                                                       beach of
of Rockanje    in  August  2013, which   he then  donated  to the   first author. This well preserved
Rockanje in August 2013, which he then donated to the first author. This well preserved tusk is straighttusk is
straight  with    no  curvation  (Figure  7). It also shows   the   characteristic groove   indicating
with no curvation (Figure 7). It also shows the characteristic groove indicating that it originated from
                                                                                                        that it
originated from the left upper jaw. Most of this gray-brown to yellow-brown specimen is covered
the left upper jaw. Most of this gray-brown to yellow-brown specimen is covered with a wafer-thin
with a wafer-thin layer of enamel. When compared to the specimens described by Machenko [15], it
layer of enamel. When compared to the specimens described by Machenko [15], it can be concluded
can be concluded that the animal was only a few months old when it died. The maximum length is
that the animal was only a few months old when it died. The maximum length is 49 mm. The depth of
49 mm. The depth of the pulp cavity measures 24 mm. Posterior of the pulp cavity, the circumference
the pulp cavity measures 24 mm. Posterior of the pulp cavity, the circumference of the tusk is 48 mm.
of the tusk is 48 mm. Measured from the back to the front, the diameter here is 17 mm, and measures
Measured from the back to the front, the diameter here is 17 mm, and measures 12 mm from medial
12 mm from medial to lateral.
to lateral.

     Figure 7. Left permanent tusk of Mammuthus primigenius. Locality: Beach of Rockanje. Collection:
     Figure 7. Left permanent tusk of Mammuthus primigenius. Locality: Beach of Rockanje. Collection: Dick
     Dick Mol (Hoofddorp). Scale bar: 5 cm. Photo: Hans Wildschut. (A) Lateral view; (B) medial view;
     Mol (Hoofddorp). Scale bar: 5 cm. Photo: Hans Wildschut. (A) Lateral view; (B) medial view; (C)
     (C) cranial view; and (D) circumference pulp cavity.
     cranial view; and (D) circumference pulp cavity.

3.8. Collection of Willy van Wingerden
3.8. Collection of Willy van Wingerden
      On 24 February 2017, Willy van Wingerden from Honselersdijk picked up a permanent tusk of
      On 24 February 2017, Willy van Wingerden from Honselersdijk picked up a permanent tusk of
a woolly mammoth from the beach of De Zandmotor. The shape of the tusk is straight (Figure 8).
a woolly mammoth from the beach of De Zandmotor. The shape of the tusk is straight (Figure 8). The
The tip is covered with a thin layer of enamel and the transition from the enamel to the dentin is
tip is covered with a thin layer of enamel and the transition from the enamel to the dentin is clearly
clearly visible. The previously described groove located on the medial side of the tusk facilitates
visible. The previously described groove located on the medial side of the tusk facilitates the
the determination as a left permanent tusk. The tusk is tan colored and the enamel tends to a dark
determination as a left permanent tusk. The tusk is tan colored and the enamel tends to a dark brown
brown color. Resembling the specimen from Dick Mol’s collection found on the beach of Rockanje
color. Resembling the specimen from Dick Mol’s collection found on the beach of Rockanje (Item 3.7),
(Item 3.7), this tusk must have belonged to a juvenile Mammuthus primigenius just a couple of months
this tusk must have belonged to a juvenile Mammuthus primigenius just a couple of months old. The
old. The maximum length is 53 mm and the pulp cavity is 21 mm deep.
maximum length is 53 mm and the pulp cavity is 21 mm deep.
Deciduous Tusks and Small Permanent Tusks of the Woolly Mammoth, Mammuthus primigenius (Blumenbach, 1799) Found on Beaches in The Netherlands - MDPI
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       Figure  8.8.Left
          Figure    Leftpermanent
                         permanenttusk    of Mammuthus
                                     tusk of              primigenius.Locality:
                                             Mammuthus primigenius.     Locality:Beach
                                                                                  Beachofof
                                                                                         dede  Zandmotor.
                                                                                            Zandmotor.      Collection:
                                                                                                         Collection:
           Figure
       Willy  van  8. Left permanent
                    Wingerden         tusk of Mammuthus
                                  (Honselersdijk).   Scale primigenius.
                                                           bar:  5 cm.  Locality:
                                                                        Photo:    Beach
                                                                                 Hans   of de Zandmotor.
                                                                                        Wildschut.  (A)   Collection:
                                                                                                        Lateral
          Willy van Wingerden (Honselersdijk). Scale bar: 5 cm. Photo: Hans Wildschut. (A) Lateral view; (B)     view;
           Willy
       (B)medial
           medialvan   Wingerden
                    view;   (C)     (Honselersdijk).
                                cranial view; and (D)Scale bar: 5 cm.
                                                       circumference  Photo:
                                                                        pulp
                 view; (C) cranial view; and (D) circumference pulp cavity.   Hans
                                                                             cavity.Wildschut.  (A) Lateral view; (B)
           medial view; (C) cranial view; and (D) circumference pulp cavity.
   3.9.
3.9.     CollectionofofMartien
      Collection              MartienLelijLelij
     3.9. Collection of Martien Lelij
        AA      superbspecimen
           superb            specimenof    ofaapermanent
                                                  permanent tusk of       of aa woolly
                                                                                 woollymammoth
                                                                                            mammothwas           wasfound
                                                                                                                       found   bybythethelate    Martien
                                                                                                                                              late  Martien
   Lelij     A superb
           from       Hoek    specimen
                                van  Holland of aon permanent        tusk ofina March
                                                                                  woolly2012.
                                                                                            mammoth             was from
                                                                                                                      foundthe  byupper
                                                                                                                                     the late     Martien
Lelij  from       Hoek      van   Holland      on thethe     local
                                                         local       beach
                                                                  beach   in March 2012.         It Itis isa tusk
                                                                                                             a tusk from the     upper right    right  jawand
                                                                                                                                                     jaw
     Lelijitfrom
   and         is      Hoek curved
                   slightly      van Holland
                                           (Figure  on9).
                                                        theItlocal    beach
                                                                clearly  has   in March
                                                                                the         2012. It isgroove
                                                                                     characteristic            a tusk onfromthe the   upperside.
                                                                                                                                  medial         rightThe
                                                                                                                                                        jaw
it is slightly curved (Figure 9). It clearly has the characteristic groove on the medial side. The brown
     and    it   is  slightly     curved     (Figure     9).  It  clearly  has   the   characteristic          groove    on   the   medial       side.  The
to brown
    dark brown to darkenamel brown layer
                                       enamel  is layer   is substantial
                                                   substantial               in thickness.
                                                                     in thickness.              The cementum
                                                                                        The cementum               layerlayer
                                                                                                                           andand the the      transition
                                                                                                                                        transition       from
     brown
   from     the  tocementum
                      dark brown     to enamel
                                         the        layer
                                               dentin    areis clearly
                                                               substantial     in thickness.
                                                                         visible.   Because      Thetip
                                                                                                the        cementum
                                                                                                               of this    layer
                                                                                                                        tusk   is  and
                                                                                                                                   not   the polished
                                                                                                                                         yet    transition
thefrom
      cementumthe          to
                      cementum the  dentin
                                       to the   are
                                                dentinclearly     visible.
                                                          are clearly        Because
                                                                          visible.         the
                                                                                     Because    tiptheofof   this
                                                                                                           tipthe   tusk
                                                                                                                ofliving   is
                                                                                                                    this tusk  not   yet
                                                                                                                                 is not      polished
                                                                                                                                            yetitpolished due
   due
to use,   to   use,    it is likely   that  it was    completely       hidden    in the   alveolus                         animal.     If   so,    would
     due it
   imply
                is
            tothat
                 use,likely
                       this
                               that itthat
                        it istusk
                               likely     wasit completely
                                    belonged    wasto acompletely
                                                                   hidden
                                                         woolly mammoth hiddenin the
                                                                                  inthatalveolus
                                                                                      the  alveolus
                                                                                           must
                                                                                                       ofofthe
                                                                                                    have       theliving
                                                                                                               died living
                                                                                                                      within
                                                                                                                            animal.
                                                                                                                            animal.
                                                                                                                                the firstIfIfso,
                                                                                                                                              so,ititwould
                                                                                                                                              year
                                                                                                                                                       would
                                                                                                                                                     of its
implyimplythat      this   tusk    belonged       to  a  woolly      mammoth         that   must       have     died    within     the first year ofofitsits
                                                                                                                                         first    year
   life.  Thisthat      this tusk belonged
                   undamaged           tusk has atolength
                                                        a woolly      mammoth
                                                                  of 102  mm. At that       mustofhave
                                                                                      the end           the pulpdied cavity,
                                                                                                                       within thethe circumference
life.life.
       This undamaged                 tusk   hashasa alength      of 102
                                                                      102mm.      At
                                                                                   At the
                                                                                        theisend    of the the pulp
                                                                                                                 pulpcavity,
                                                                                                                         cavity,the the circumference
   of theThis tuskundamaged
                       is 55 mm. The     tuskdiameter   length
                                                           of theofoval    mm. cavity
                                                                           pulp              end
                                                                                               19 ofmm        at maximum.         The circumference
                                                                                                                                        enamel layer
of the
     of   tusk
         the    tuskis 55is  mm.
                            55   mm.The Thediameter
                                               diameter   ofofthetheoval
                                                                      oval pulp
                                                                            pulp   cavity
                                                                                    cavity   is
                                                                                             is 19
                                                                                                 19    mm
                                                                                                       mm      at maximum.
                                                                                                               at  maximum.The       Theenamel
                                                                                                                                             enamellayer layer
   measures from the tip of the tusk to the rear over a maximum length of 66 mm.
measures          from     the   tip  of  the  tusk    to  the   rear  over
     measures from the tip of the tusk to the rear over a maximum length       a maximum          length        of 66  mm.
                                                                                                                       mm.

         Figure 9. Right permanent tusk of Mammuthus primigenius. Locality: Beach of Hoek van Holland.
           Figure
       Figure      9. Right
               9. Right
         Collection:        permanent
                         permanent
                      Martien            tusk
                                       tusk
                              Lelij (Hoek   ofofMammuthus
                                            van   Mammuthus
                                                 Holland).     primigenius.
                                                           Scale  bar: 5 cm.Locality:
                                                             primigenius.   Locality: Beach
                                                                                       Beach
                                                                             Photo: Hans    ofofHoek
                                                                                                 Hoekvan
                                                                                           Wildschut.  vanHolland.
                                                                                                      (A)   Holland.
                                                                                                           Lateral
           Collection: Martien
         view; (B) Martien
       Collection:             Lelij
                    medial view;      (Hoek
                                  (C) cranial
                            Lelij (Hoek       van Holland).
                                          van view; and (D)
                                               Holland).    Scale  bar: 5
                                                            circumference
                                                         Scale            cm. Photo:  Hans
                                                                            pulp cavity.
                                                               bar: 5 cm. Photo:            Wildschut.  (A) Lateral
                                                                                  Hans Wildschut. (A) Lateral  view;
           view; (B) medial view; (C) cranial view; and (D) circumference pulp cavity.
       (B) medial view; (C) cranial view; and (D) circumference pulp cavity.
Deciduous Tusks and Small Permanent Tusks of the Woolly Mammoth, Mammuthus primigenius (Blumenbach, 1799) Found on Beaches in The Netherlands - MDPI
Quaternary
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           2018, 2,
                 1, x7 FOR PEER REVIEW                                                                                        10
                                                                                                                              10 of
                                                                                                                                 of 14
                                                                                                                                    14

3.10. Collection of Trudy Langeveld
3.10. Collection of Trudy Langeveld
      A right permanent juvenile tusk was collected by Trudy Langeveld from Voorhout in January
      A right
2011 on           permanent
           the beach    of Hoek  juvenile  tusk was
                                   van Holland.     Thecollected     by Trudy
                                                          sand originated         Langeveld
                                                                               from                from Voorhout
                                                                                        the extraction     areas to the  in north
                                                                                                                             Januaryof
2011   on  the   beach  of Hoek     van  Holland.   The   sand    originated    from     the  extraction
the Euro-Maasgeul, off the coast of South Holland. It has been added to her collection under catalog       areas   to  the   north  of
the Euro-Maasgeul,
number     141.            off the coast of South Holland. It has been added to her collection under catalog
numberThis141.
             larger specimen, mostly yellow-brown colored, is slightly damaged; the tip has been broken
      Thispossibly
recently,     larger specimen,
                        due to the mostly   yellow-brown
                                       suction  action of the  colored,   is slightlymachine
                                                                   sand dredging          damaged;(Figure
                                                                                                       the tip10).
                                                                                                                has It been    broken
                                                                                                                          is slightly
recently, possibly
rounded      medially,due     to thethe
                         showing       suction  action of groove.
                                          characteristic     the sandDeposits
                                                                         dredging     ofmachine
                                                                                          wafer-thin (Figure
                                                                                                         black10).
                                                                                                                 enamelIt is are
                                                                                                                             slightly
                                                                                                                                  still
roundedonmedially,
present                  showing
                the lateral side of the
                                      thischaracteristic
                                           tusk. The marks   groove.
                                                                 of thisDeposits
                                                                          permanent   of wafer-thin
                                                                                           tusk suggest  black
                                                                                                             thatenamel       are still
                                                                                                                   it had already
present on
emerged         the lateral
             from   the tuskside   of this
                                socket  andtusk.
                                              mustThe   marks
                                                     have    been ofvisible
                                                                     this permanent
                                                                             in the living  tuskanimal.
                                                                                                  suggestWhenthat itcompared
                                                                                                                       had already  to
emerged
the  specimenfromdescribed
                    the tusk socket
                                by Vananddermust   have
                                              Plicht       been
                                                      et al. ([10]visible  in the2)
                                                                    see Figure      living
                                                                                       it cananimal.    When compared
                                                                                               be concluded                     to the
                                                                                                                  that the animal
specimen
was   12–24described
                 months byoldVan     der Plicht
                                 at time         et al. ([10]
                                           of death.           see Figure has
                                                        The specimen         2) it acan   be concluded
                                                                                       preserved      lengththatofthe102animal
                                                                                                                           mm, wasbut
12–24 months old
reconstructing      theat time
                        total     of death.
                               length   usingThe
                                               thespecimen
                                                   specimenhas      a preserved
                                                                 from  the collectionlength of of 102 mm,
                                                                                               Martien        but
                                                                                                          Lelij  forreconstructing
                                                                                                                      comparison,
theleast
at   total30length
              mm isusing    the specimen
                      missing.    The depthfrom
                                              of thethe  collection
                                                       pulp           of84
                                                              cavity is   Martien
                                                                             mm and    Lelij
                                                                                          its for comparison,
                                                                                               maximum             at least
                                                                                                            diameter           30 mm
                                                                                                                           measures
is missing.
28 mm.          The  depth   of  the  pulp  cavity  is 84  mm    and  its maximum          diameter     measures      28  mm.

      Figure
      Figure 10.
              10. Right
                   Right permanent
                         permanent tusk
                                      tusk of
                                           of Mammuthus
                                              Mammuthus primigenius.
                                                           primigenius. Locality:
                                                                         Locality: Beach
                                                                                    Beachof
                                                                                          ofHoek
                                                                                            Hoek van
                                                                                                  van Holland.
                                                                                                      Holland.
      Collection:
      Collection: Trudy Langeveld (Voorhout). Scale bar: 5 cm. Photo: Hans Wildschut. (A) Medial view;
                   Trudy  Langeveld    (Voorhout). Scale bar: 5 cm. Photo:  Hans   Wildschut. (A) Medial view;
      (B)
      (B) lateral
          lateral view;
                  view; (C)
                        (C) cranial
                            cranial view;
                                    view; and
                                          and (D)
                                               (D) circumference  pulp cavity.
                                                   circumference pulp   cavity.

3.11.
3.11. Collection
      Collection of
                 of Henk
                    Henk Mulder
                         Mulder
     In
     In November
        November 2017,
                    2017, Henk
                           Henk Mulder
                                  Mulder from
                                           from Monster
                                                  Monster picked
                                                           picked up
                                                                  up aa left
                                                                         left permanent
                                                                              permanent tusktusk of
                                                                                                 of aa young
                                                                                                       young
woolly  mammoth    from   the beach  of De  Zandmotor.   Unfortunately,   the  tip of the  tusk
woolly mammoth from the beach of De Zandmotor. Unfortunately, the tip of the tusk was broken    was   broken
                                                                                                          due
due  to mechanical
to mechanical        action.
               action.        However,
                       However,           thecavity
                                   the pulp    pulp was
                                                     cavity wasThis
                                                         intact. intact.
                                                                     tuskThis
                                                                            was tusk
                                                                                 fromwas     fromindividual
                                                                                        an older    an older
individual than thefrom
than the specimen   specimen   from theofcollection
                         the collection             of Trudy(Item
                                           Trudy Langeveld    Langeveld   (Item
                                                                  3.10). The     3.10).isThe
                                                                               length        length
                                                                                          about      is about
                                                                                                150 mm    and
150 mm  and  the depth  of the pulp  cavity
the depth of the pulp cavity is 55 mm.      is 55 mm.

4.
4. The
   The Juvenile
       Juvenile Tusk Look-Alike
                Tusk Look-Alike
     The
     The preceding
         preceding text
                    text gives
                         gives an
                               an overview
                                  overview of
                                            of known
                                               known deciduous
                                                      deciduous tusks
                                                                    tusks and
                                                                          and permanent
                                                                               permanent tusks
                                                                                            tusks of
                                                                                                   of very
                                                                                                      very
young
young woolly
       woolly mammoths.
              mammoths. TheThe knowledge
                                knowledge is
                                          is valuable
                                             valuable as
                                                      as aa reference
                                                            reference collection,
                                                                      collection, and
                                                                                  and also
                                                                                      also to
                                                                                           to distinguish
                                                                                              distinguish
them from confounding elements that look similar but yet have a different origin; in particular, the
following dental element.
Quaternary 2018, 1, 7                                                                                                      11 of 14

them    from
 Quaternary     confounding
             2018,  2, x FOR PEERelements
                                     REVIEW that look similar but yet have a different origin; in particular,                11 of 14
the following dental element.
       Erwinvan
      Erwin      vander derLee
                             Leefound
                                  foundaafragment
                                              fragmentof   ofaadental
                                                                 dentalelement
                                                                         elementin  inthe
                                                                                       thesupplemented
                                                                                            supplementedsand  sandof ofthe
                                                                                                                         thebeach
                                                                                                                             beach
 ofHoek
of  Hoekvan vanHolland
                   Hollandin   inAugust
                                   August2017  2017(Figure
                                                      (Figure11).
                                                                11).The
                                                                     Thespecimen
                                                                          specimenhas  hassome
                                                                                             somecharacteristics
                                                                                                    characteristicsof ofaatusk
                                                                                                                            tuskof of
aajuvenile
   juvenilemammoth.
                mammoth. It    It is
                                   is fragile,
                                       fragile, not
                                                  not fully
                                                       fully developed,
                                                              developed, without a root and a very      very deep
                                                                                                             deep pulp cavity.
                                                                                                                            cavity.
 Themaximum
The    maximumlength   lengthisis4646mm.mm.ItItappears
                                                 appearsto  tobebecomplete,
                                                                    complete,from
                                                                                fromthethepulp
                                                                                           pulpcavity
                                                                                                   cavityon
                                                                                                          onthe
                                                                                                              theposterior
                                                                                                                   posteriorside
                                                                                                                               side
 tothe
to  thetip
         tipononthe
                  theanterior
                         anteriorside.
                                    side.ItItisisslightly
                                                  slightlycurved
                                                            curvedand andstrongly
                                                                           stronglyflattened
                                                                                      flattenedlaterally.
                                                                                                  laterally.The
                                                                                                             Thetiptipisiscovered
                                                                                                                           covered
 by  a thin  cap.   The    circumference        of this  tusk  is 28 mm    at the  pulp  cavity   and
by a thin cap. The circumference of this tusk is 28 mm at the pulp cavity and 26 mm in the middle of   26 mm    in the   middle    of
 thetusk.
the   tusk.The
             Thedepth
                    depthof  ofthis
                                thiscavity
                                        cavityappears
                                                 appearsto  tobebe22
                                                                   22mm.
                                                                       mm.The Thespecimen
                                                                                   specimenconsists
                                                                                                consistsofofdentin
                                                                                                             dentincompletely
                                                                                                                      completely
 coveredwith
covered     withaafilmfilmofofenamel,
                               enamel,whichwhichin   inturn
                                                        turnisiscovered
                                                                 coveredby byaathin
                                                                                 thinlayer
                                                                                      layerof ofcementum.
                                                                                                 cementum.The  Thecross-section
                                                                                                                    cross-section
 ofthe
of  thedental
         dentalelement
                   elementisisflattened,
                                 flattened,giving
                                                givingan anoval
                                                             ovalshape
                                                                   shapeover
                                                                          overthetheentire
                                                                                     entirelength.
                                                                                              length.The
                                                                                                      Themaximum
                                                                                                           maximumdiameterdiameter
 ofthe
of  thetusk,
          tusk,measured
                 measuredfrom   fromfrontfronttotoback
                                                     backisis55mm,
                                                                 mm,ininanatomical
                                                                          anatomicalposition.
                                                                                         position.ItItmeasures
                                                                                                        measures10  10mm mmfrom
                                                                                                                              from
 medialto
medial     tolateral.
               lateral.TheThepulp
                               pulpcavity
                                        cavityisisextremely
                                                    extremely small
                                                                  small compared
                                                                         compared to  to all
                                                                                         all known
                                                                                               known small
                                                                                                       small permanent
                                                                                                              permanent tusks.
                                                                                                                             tusks.
 Moreover,this
Moreover,       thisspecimen
                        specimenisistoo  tooflattened,
                                               flattened,seen
                                                            seenfrom
                                                                   fromfront
                                                                          fronttotoback.
                                                                                     back. These
                                                                                             These are
                                                                                                     arenot
                                                                                                         notcharacteristics
                                                                                                               characteristicsof   of
 newly-developed permanent
newly-developed            permanent tusks.tusks. Furthermore, the morphology of the find             find is
                                                                                                            is too
                                                                                                                too irregular
                                                                                                                     irregular in  in
 structure    to  be  determined        as a (permanent)
structure to be determined as a (permanent) tusk.              tusk.

       Figure11.
      Figure   11.Digitelle
                   Digitelleof
                             ofaapremolar
                                   premolarofofMammuthus      primigenius. Locality:
                                                Mammuthusprimigenius.      Locality: Beach
                                                                                     Beach of
                                                                                            of Hoek
                                                                                               Hoek van
                                                                                                     van Holland.
                                                                                                          Holland.
       Collection: Erwin   van  der Lee  (Rotterdam).  Photo: Hans  Wildschut.  (A)  Caudal view; (B) cranial
      Collection: Erwin van der Lee (Rotterdam). Photo: Hans Wildschut. (A) Caudal view; (B) cranial view;    view;
       (C)medial
      (C) medialview;
                   view;(D)
                          (D)lateral
                               lateralview;
                                       view;and
                                             and(E)
                                                 (E)circumference
                                                     circumferencepulp
                                                                     pulpcavity.
                                                                           cavity.

       This specimen must be identified as one of the digitelles, which is a part of lamella of a mammoth
      This specimen must be identified as one of the digitelles, which is a part of lamella of a mammoth
 molar. A pillar or a slat shows deep incisions. It appears that such a pillar is composed of several
molar. A pillar or a slat shows deep incisions. It appears that such a pillar is composed of several
 adjacent, and, on the bottom, closely connected digits that are flattened from the front to the rear.
adjacent, and, on the bottom, closely connected digits that are flattened from the front to the rear. Such a
 Such a pillar is made up of several varying digitelles. Several digits and the pillars form a plate. The
pillar is made up of several varying digitelles. Several digits and the pillars form a plate. The digits
 digits are usually connected at the base. This is not always the case with premolars. The size of this
are usually connected at the base. This is not always the case with premolars. The size of this digitelle
 digitelle suggests that this must have been part of a third or fourth premolar.
suggests that this must have been part of a third or fourth premolar.
 5. Conclusions
      The tusks of juvenile Mammuthus primigenius discussed in this paper are of great scientific value as
 they are still very rare in the fossil record. Because of sand supplementation activities of the Dutch
 beaches, deciduous tusks and very small permanent tusks are now occasionally found. These elements
 are very important sources for mammalian paleontological research. However, the chance of recognition
Quaternary 2018, 1, 7                                                                                      12 of 14

5. Conclusions
     The tusks of juvenile Mammuthus primigenius discussed in this paper are of great scientific
value as they are still very rare in the fossil record. Because of sand supplementation activities
of the Dutch beaches, deciduous tusks and very small permanent tusks are now occasionally found.
These elements are very important sources for mammalian paleontological research. However,
the chance of recognition is also slim. The tusks described and illustrated here can aid in recognition
of these particular mammoth remains and increase the likelihood that they will be preserved for the
fossil record.

5.1. Basic Characteristics of Deciduous Tusks
     A deciduous tusk is somewhat club shaped and has a relatively large root and a well-developed
cap on the tip, which is covered with a thin enamel layer that in turn is covered by a thin layer of
cementum. It may measure around 35–100 mm in length and usually nearly straight or only slightly
nonlinear. The cross-section is more or less oval shaped—the lateral side being more curved than the
medial side. A shallow groove may be visible on the medial side of the tusk (Figures 3A, 4A, 5A, 6B,
7B, 8B, 9B and 10A).

5.2. Basic Characteristics of Juvenile Permanent Tusks
     The major distinction between the deciduous and permanent tusk is the relatively deep pulp
cavity, which may measure up to about half of its total length. Permanent tusks do not have a root.
The length of the smallest known specimen measures 37 mm and it is covered with a very thin layer of
enamel. The common features with deciduous tusks are the cross-section, the lateral side being more
curved compared to the medial side. The medial side of the tusk is characterized by a shallow groove.
However, a correct identification of fossil remains requires careful comparisons. To determine whether
the specimen is a deciduous tusk or a newly-developed permanent tusk, a thorough knowledge is
required of the tusk and the development of proboscideans. It was only after extensive comparisons
that the discovery of Erwin van der Lee could be definitively classified as a digitelle of a lamella or
plate of a Mammuthus primigenius, rather than a juvenile tusk.
     Tusks of proboscideans have a complex structure with chevron shaped lines (Schreger lines) and
growth rings much like trees, which carry the complete life history. A detailed analysis of a tusk’s
interior would provide more information about the first years of life of mammoths which inhabited
the vast plains between the British Isles and the continent of Europe in the Late Pleistocene. Then,
we might be able to judge the rate at which newborn mammoth calves have grown, and whether or
not that matches what we know about elephants in Africa and Asia. Hopefully, more such finds will
be made in sufficient quantities to sacrifice one or more of good quality to be subjected to the required
destructive tests for that.

Author Contributions: The authors all contributed to the determinations, to the analysis of the results and to the
writing of the manuscript.
Funding: No funding was received for the production of this paper.
Acknowledgments: The authors would like to express their gratitude to Erwin van der Lee from Rotterdam for
making available his discovery of the digitelle of a premolar woolly mammoth for identification and research.
Once more, we would like to thank Jan van der Steeg in Losser and Walter Langendoen in Hellevoetsluis,
for making available the deciduous tusks in their care; Peter Soeters in Delft for his donation of a number
of mammoth finds, including the very small permanent tusk; the late Martien Lelij from Hoek of Holland,
Hansjorg Ahrens from Leiden, Trudy Langeveld from Voorhout, Henk Mulder from Monster, Donny Chrispijn
from Hoek van Holland and Willy van Wingerden from Honselersdijk for providing their exceptional finds of
very small permanent tusks for description; Hans Wildschut, Hoofddorp, for the photography; and Klaas Post
and Bram Langeveld (Natuurhistorisch Museum Rotterdam) for reviewing earlier drafts of this paper.
Conflicts of Interest: The authors declare no conflict of interest.
Ethical Statement: In The Netherlands, collecting fossils from public places is legal. However, fossil collection
elsewhere may be limited or prohibited by law. Also none of the authors are a member of any professional
Quaternary 2018, 1, 7                                                                                               13 of 14

society with a code of ethics that recommends against publishing work on specimens that are held in private
collections. Such a code would be counterproductive in The Netherlands. The vast public beaches contain an
abundance of small fossil remains. Over the years, the number of non-professional paleontologists has increased
steadily, and they may have collected many rare specimens that would not have been discovered otherwise.
Non-professionals are a valuable resource to the field of paleontology here. In the Netherlands we are facing a
unique situation. For almost four decades, professionals and non-professionals have been united in the active
Werkgroep Pleistocene Zoogdieren (working group Pleistocene Mammals, www.pleistocenemammals.com),
which organizes very well attended fossil determination sessions several times a year for their ever-increasing
collections. At these meetings, non-professionals can learn and have their fossil finds identified by professionals.
The professional scientific community benefits from these meetings in getting an overview of all the discoveries
made in the field. Knowledge and information is shared mutually. In this way, many rare specimens have emerged,
enriching the Pleistocene fauna of the North Sea. Often these collectors develop an interest in the whereabouts of
their finds and many educate themselves to become ‘citizen scientists’. In addition, the collectors are encouraged
to donate or loan their specimens to museums for availability for research. Moreover, replicas are routinely
produced for display. Most collectors are happy to have their specimens examined by scholars. Publicizing about
these noteworthy finds in leading international media is a logical part of this process.

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