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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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
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.
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
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
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.
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.
Quaternary 2018, 1, 7 7 of 14 Quaternary 2018, 2, x FOR PEER REVIEW 7 of 14 Quaternary 2018, 2, x FOR PEER REVIEW 7 of 14 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.
Quaternary 2018, 1, 7 8 of 14 Quaternary 2018, 2, x FOR PEER REVIEW 8 of 14 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.
Quaternary 2018, 1, 7 9 of 14 Quaternary 2018, 2, x FOR PEER REVIEW 9 of 14 Quaternary 2018, 2, x FOR PEER REVIEW 9 of 14 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.
Quaternary Quaternary 2018, 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. References 1. Mol, D.; de Vos, J.; Bakker, R.; van Geel, B.; Glimmerveen, J.; van der Plicht, J.; Post, K. Kleine Encyclopedie Van Het Leven in Het Pleistoceen—Mammoeten, Neushoorns en Andere Dieren van de Noordzeebodem; Uitgeverij Veen Magazines B.V.: Diemen, The Netherlands, 2008; ISBN 9789085710981. 2. Van Kolfschoten, T.; Laban, C. Pleistocene terrestrial mammal faunas from the North Sea. Meded. Rijks Geol. Dienst 1995, 52, 135–151. 3. Mol, D. Twee etmalen korren in de Eurogeul: trok de kotter OD7 door een mammoetkerkhof? Afzettingen 2012, 33, 7–10. 4. Mol, D.; Post, K. Gericht korren op de Noordzee voor de zoogdierpaleontologie: Een historisch overzicht van de uitgevoerde expedities. Cranium 2010, 2, 14–28. 5. Laban, C.; Rijsdijk, K.F. De Rijn-Maasdelta’s in de Noordzee. Grondboor Hamer 2002, 56, 60–65. 6. Rijsdijk, K.F.; Kroon, I.C.; Meijer, T.; Passchier, S.; van Dijk, T.A.G.P.; Bunnik, F.P.M.; Janse, A.C. Reconstructing Quaternary Rhine–Meuse dynamics in the southern North Sea: Architecture, seismolithofacies associations and malacological biozonation. J. Quat. Sci. 2013, 28, 453–466. [CrossRef] 7. Post, K. A Weichselian marine mammal assemblage from the southern North Sea. Deinsea 2005, 11, 21–27. 8. Mol, D.; Borst, W.; Reumer, J. De eerste fossiele hyenakeutel uit de Noordzee. Straatgras 2010, 22, 91–93. 9. Mol, D.; Langeveld, B.; Janse, A.; Langendoen, W.; Smolarz, J. Determinatiedag fossiele strandvondsten van Maasvlakte 2 in Futureland: Een verslag. Cranium 2015, 32, 49–58. 10. Van der Plicht, J.; Post, K.; Mol, D. Over aasvliegen en een mammoetkalf uit de Eurogeul. Cranium 2012, 29, 14–19. 11. Dieleman, F. Overzicht van strandvondsten van woelmuizen en andere kleine zoogdieren langs de Nederlandse stranden: Stand van zaken 2013. Afzettingen 2013, 34, 144–172. 12. Langeveld, B. Trogontherium cuvieri Fischer (Castoridae) van het strand van Hoek van Holland en de Zandmotor. Cranium 2013, 30, 8–12. 13. Langeveld, B. Fossiele omhulsels van poppen van de subarctische aasvlieg Protophormia terraenovae Robineau-Desvoidy, 1830 voor het eerst van opgespoten stranden (Eurogeulgebied). Cranium 2015, 32, 34–38. 14. Langeveld, B. Vondsten van de reuzenalk Pinguinus impennis (Linnaeus, 1758) (Aves) uit het Eurogeulgebied. Cranium 2015, 32, 19–27. 15. Maschenko, E.N. Individual development, biology and evolution of the woolly mammoth. Cranium 2002, 19, 4–120. 16. Maschenko, E.N.; Boeskorov, G.G.; Baranov, V.A. Morphology of a Mammoth Calf (Mammuthus prinigenius) from Ol’chan (Oimiakon, Yakutia). Paleontol. J. 2013, 47, 425–438. [CrossRef] 17. Rountrey, A.N.; Fisher, D.C.; Tikhonov, A.N.; Kosintsev, P.A.; Lazarev, P.A.; Boeskorov, G.; Buigues, B. Early tooth development, gestation, and season of birth in mammoths. Quat. Int. 2012, 255, 196–205. [CrossRef] 18. Mol, D. Mammoth fossils recovered from the seabed between the British Isles and the European continent. Bulletin de Musée d’Anthropologie préhistorique de Monaco 2016, 6, 129–142. 19. Maschenko, E.; Tikhonov, A.N.; MacPhee, R.D. Mammoth calf from Bolshoi Lyakhovskii Island (New Siberian Islands, Arctic Siberia). Russ. J. Theriol. 2005, 4, 79–88. [CrossRef]
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