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Smithsonian at the Poles: Contributions to International Polar

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Consider<strong>at</strong>ions of An<strong>at</strong>omy,<br />

Morphology, Evolution, and<br />

Function for Narwhal Dentition<br />

Martin T. Nweeia, Frederick C. Eichmiller,<br />

Cornelius Nutarak, Naomi Eidelman,<br />

Anthony A. Giuseppetti, Janet Quinn, James<br />

G. Mead, Kaviqanguak K’issuk, Peter V.<br />

Hauschka, Ethan M. Tyler, Charles Potter,<br />

Jack R. Orr, Rasmus Avike, Pavia Nielsen,<br />

and David Angn<strong>at</strong>siak<br />

Martin T. Nweeia, Harvard University, School of<br />

Dental Medicine, Bos<strong>to</strong>n, MA, USA. Cornelius<br />

Nutarak and David Angn<strong>at</strong>siak, Community<br />

of Mittim<strong>at</strong>ilik, Nunavut, Canada. Frederick C.<br />

Eichmiller, Delta Dental of Wisconsin, Stevens<br />

Point, WI, USA. Naomi Eidelman, Anthony A.<br />

Giuseppetti, and Janet Quinn, ADAF Paffenbarger<br />

Research Center, N<strong>at</strong>ional Institute of Standards<br />

and Technology, Gai<strong>the</strong>rsburg, MD, USA.<br />

James G. Mead and Charles Potter, <strong>Smithsonian</strong><br />

Institution, Division of Mammals, Department of<br />

Zoology, Washing<strong>to</strong>n, DC, USA. Kaviqanguak<br />

K’issuk and Rasmus Avike, Community of Qaanaaq,<br />

Greenland. Peter V. Hauschka, <strong>Smithsonian</strong><br />

Institution, Division of Mammals, Department of<br />

Zoology, Washing<strong>to</strong>n, DC, USA. Ethan M. Tyler,<br />

N<strong>at</strong>ional Institutes of Health, Clinical Center,<br />

Be<strong>the</strong>sda, MD, USA. Jack R. Orr, Fisheries and<br />

Oceans Canada, Arctic Research Division, Winnipeg,<br />

MB, Canada. Pavia Nielsen, Community of<br />

Uummannaq, Greenland. Corresponding author:<br />

M. T. Nweeia (martin_nweeia@hsdm.harvard.<br />

edu). Accepted 19 May 2008.<br />

ABSTRACT. Interdisciplinary studies of narwhal cranial and <strong>to</strong>oth an<strong>at</strong>omy are combined<br />

with Inuit traditional knowledge <strong>to</strong> render a more complete description of <strong>to</strong>oth- rel<strong>at</strong>ed<br />

structures and <strong>to</strong> propose a new hypo<strong>the</strong>sis for tusk function in <strong>the</strong> adult male. Gross<br />

an<strong>at</strong>omy fi ndings from computed <strong>to</strong>mography (CT) and magnetic resonance (MR) imaging<br />

and dissections of an adult male and female and one fetus, four <strong>to</strong> six months in development,<br />

were documented. Computed <strong>to</strong>mography scans rendered images of <strong>the</strong> tusks and<br />

vestigial teeth and <strong>the</strong>ir shared sources of innerv<strong>at</strong>ion <strong>at</strong> <strong>the</strong> base of <strong>the</strong> tusks. Paired and<br />

asymmetrical tusks and vestigial teeth were observed in all three samples, and <strong>the</strong>ir rel<strong>at</strong>ive<br />

positions reversed during development. Vestigial teeth shifted anteriorly during growth,<br />

and <strong>the</strong> developing tusks moved posteriorly as <strong>the</strong>y developed. Examin<strong>at</strong>ion of tusk microan<strong>at</strong>omy<br />

revealed <strong>the</strong> presence of a dentinal tubule network with lumena approxim<strong>at</strong>ely<br />

2 micrometers in diameter and 10– 20 micrometers apart over <strong>the</strong> pulpal and erupted tusk<br />

surfaces. Orifi ces were present on <strong>the</strong> cementum surface indic<strong>at</strong>ing direct communic<strong>at</strong>ion<br />

and sensory capability from <strong>the</strong> environment <strong>to</strong> <strong>the</strong> inner pulpal wall. Flexural strength of<br />

95 MPa <strong>at</strong> mid tusk and 165 MPa <strong>at</strong> <strong>the</strong> base indic<strong>at</strong>ed resistance <strong>to</strong> high fl exural stresses.<br />

Inuit knowledge describes a tusk with remarkable and combined strength and fl exibility.<br />

Elder observ<strong>at</strong>ions of an<strong>at</strong>omy are described by variable phenotypes and classifi ed by skin<br />

color<strong>at</strong>ion, sex, and tusk expression. Behavioral observ<strong>at</strong>ions of males leading seasonal<br />

migr<strong>at</strong>ion groups, nonaggressive tusk encounters, and frequent sightings of smaller groups<br />

separ<strong>at</strong>ed by sex add <strong>to</strong> <strong>the</strong> discussion of tusk function.<br />

INTRODUCTION<br />

The narwhal is unique among <strong>to</strong>o<strong>the</strong>d marine mammals and exhibits unusual<br />

fe<strong>at</strong>ures, which are described in <strong>the</strong> liter<strong>at</strong>ure (Figure 1). A single 2– 3 m tusk is<br />

characteristic of adult males (Tomlin, 1967). Tusks are horizontally imbedded in

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