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<div class=Section1>

<h1 style='margin:.75pt'><span style='font-size:10.0pt;font-family:"Arial","sans-serif";
color:#5D5D5D'>Zoological Journal of the Linnean Society<o:p></o:p></span></h1>

<p class=MsoNormal style='margin-left:0in;text-indent:-.25in;mso-list:l2 level1 lfo5;
vertical-align:baseline'><![if !supportLists]><span style='font-size:10.0pt;
font-family:Symbol'><span style='mso-list:Ignore'>·<span style='font:7.0pt "Times New Roman"'>        
</span></span></span><![endif]><span class=article-headermeta-info-label><b><span
style='font-size:10.0pt;font-family:"Arial","sans-serif";border:none windowtext 1.0pt;
padding:0in'>First published:</span></b></span><span
class=apple-converted-space><b><span style='font-size:10.0pt;font-family:"Arial","sans-serif";
border:none windowtext 1.0pt;padding:0in'> </span></b></span><span
style='font-size:10.0pt;font-family:"Arial","sans-serif"'>7 September 2016 <o:p></o:p></span></p>

<p class=MsoNormal style='margin-left:0in;text-indent:-.25in;mso-list:l2 level1 lfo5;
vertical-align:baseline'><![if !supportLists]><span style='font-size:10.0pt;
font-family:Symbol'><span style='mso-list:Ignore'>·<span style='font:7.0pt "Times New Roman"'>        
</span></span></span><![endif]><span class=article-headermeta-info-label><b><span
style='font-size:10.0pt;font-family:"Arial","sans-serif";border:none windowtext 1.0pt;
padding:0in'>DOI:</span></b></span><span class=apple-converted-space><b><span
style='font-size:10.0pt;font-family:"Arial","sans-serif";border:none windowtext 1.0pt;
padding:0in'> </span></b></span><span class=article-headermeta-info-data><span
style='font-size:10.0pt;font-family:"Arial","sans-serif";border:none windowtext 1.0pt;
padding:0in'>10.1111/zoj.12456</span></span><span style='font-size:10.0pt;
font-family:"Arial","sans-serif"'> <o:p></o:p></span></p>

<h1 style='mso-margin-top-alt:0in;margin-right:0in;margin-bottom:5.05pt;
margin-left:0in;vertical-align:baseline'><span style='font-size:10.0pt;
font-family:"Arial","sans-serif";color:black'><o:p> </o:p></span></h1>

<h1 style='mso-margin-top-alt:0in;margin-right:0in;margin-bottom:5.05pt;
margin-left:0in;vertical-align:baseline'><span style='font-size:19.0pt;
font-family:"Arial","sans-serif";color:black'>Macroraptorial sperm whales
(Cetacea, Odontoceti, Physeteroidea) from the Miocene of Peru<o:p></o:p></span></h1>

<h3 style='margin:0in;margin-bottom:.0001pt;text-indent:-.25in;mso-list:l1 level1 lfo4;
background:white;vertical-align:baseline'><![if !supportLists]><span
style='font-size:10.0pt;font-family:Symbol;font-weight:normal'><span
style='mso-list:Ignore'>·<span style='font:7.0pt "Times New Roman"'>        
</span></span></span><![endif]><span style='font-size:12.5pt;font-family:"Arial","sans-serif";
font-weight:normal'>Olivier Lambert, </span><span style='font-size:12.5pt;
font-family:"Arial","sans-serif";font-weight:normal'>Giovanni Bianucci, Christian
De Muizon<o:p></o:p></span></h3>

<p class=MsoNormal style='mso-margin-top-alt:0in;margin-right:0in;margin-bottom:
6.0pt;margin-left:0in;text-indent:-.25in;mso-list:l1 level2 lfo4;background:
white;vertical-align:baseline'><![if !supportLists]><span style='font-size:
10.0pt;font-family:"Arial","sans-serif"'><span style='mso-list:Ignore'>0.<span
style='font:7.0pt "Times New Roman"'>     </span></span></span><![endif]><span
style='font-size:10.0pt;font-family:"Arial","sans-serif"'><o:p> </o:p></span></p>

<h2 style='mso-margin-top-alt:0in;margin-right:0in;margin-bottom:.1in;
margin-left:0in;vertical-align:baseline'><span style='font-size:16.0pt;
font-family:"Arial","sans-serif"'>Abstract<o:p></o:p></span></h2>

<p style='margin:0in;margin-bottom:.0001pt;vertical-align:baseline;outline: 0px;
background-position-x:0px;background-position-y:0px'><span style='font-family:
"Arial","sans-serif"'>The three extant sperm whale species, the giant<span
class=apple-converted-space> </span><em><span style='font-family:"Arial","sans-serif";
border:none windowtext 1.0pt;padding:0in'>Physeter macrocephalus</span></em><span
class=apple-converted-space> </span>and the diminutive<em><span
style='font-family:"Arial","sans-serif";border:none windowtext 1.0pt;
padding:0in'>Kogia</span></em><span class=apple-converted-space> </span>species,
are relict members of the superfamily Physeteroidea, whose evolutionary history
is traced back to the late Oligocene. Although well-preserved and diagnostic
cranial remains are relatively scarce, the physeteroid fossil record reveals a
considerable degree of morphological disparity (especially during the Miocene),
suggesting that sperm whales occupied a broader range of ecological niches in
the past. Here, we provide detailed descriptions and a (re)analysis of several
new and established middle–late Miocene stem physeteroids from the Pisco
Formation of southern Peru. In particular, we (1) further describe the holotype
of<span class=apple-converted-space> </span><em><span style='font-family:
"Arial","sans-serif";border:none windowtext 1.0pt;padding:0in'>Acrophyseter
deinodon</span></em><span class=apple-converted-space> </span>from the
latest Tortonian–Messinian of Sud-Sacaco, with new information on
previously unpublished portions of the skull (including the ear bones) and a
discussion of its masticatory musculature; (2) diagnose a new species of <em><span
style='font-family:"Arial","sans-serif";border:none windowtext 1.0pt;
padding:0in'>Acrophyseter</span></em>,<span class=apple-converted-space><b><span
style='border:none windowtext 1.0pt;padding:0in'> </span></b></span><em><b><span
style='font-family:"Arial","sans-serif";border:none windowtext 1.0pt;
padding:0in'>A. robustus</span></b></em><span class=apple-converted-space><b><span
style='border:none windowtext 1.0pt;padding:0in'> </span></b></span><b><span
style='border:none windowtext 1.0pt;padding:0in'>sp. nov.,</span></b><span
class=apple-converted-space> </span>based on a finely preserved skull with
some associated cranial remains from the late Serravallian–Tortonian
(> 9.2 Ma) of Cerro la Bruja; (3) provisionally refer a skull from
Cerro los Quesos, which has been radiometrically dated to the Messinian
(6.9–6.7 Ma), to<span class=apple-converted-space> </span><em><span
style='font-family:"Arial","sans-serif";border:none windowtext 1.0pt;
padding:0in'>Acrophyseter</span></em><span class=apple-converted-space> </span>sp.;
and (4) further describe the skull of the giant raptorial sperm whale<span
class=apple-converted-space> </span><em><span style='font-family:"Arial","sans-serif";
border:none windowtext 1.0pt;padding:0in'>Livyatan melvillei</span></em><span
class=apple-converted-space> </span>from Cerro Colorado, recently re-dated
to the Tortonian (9.9–8.9 Ma) based on the associated diatom fauna
and Ar/Ar dating. A phylogenetic analysis based on 53 characters and 21
physeteroid species confirms the monophyly of<span class=apple-converted-space> </span><em><span
style='font-family:"Arial","sans-serif";border:none windowtext 1.0pt;
padding:0in'>Acrophyseter</span></em><span class=apple-converted-space> </span>and
groups this genus with the larger, middle to late Miocene macroraptorial stem
physeteroids<span class=apple-converted-space> </span><em><span
style='font-family:"Arial","sans-serif";border:none windowtext 1.0pt;
padding:0in'>Brygmophyseter</span></em><span class=apple-converted-space> </span>and<span
class=apple-converted-space> </span><em><span style='font-family:"Arial","sans-serif";
border:none windowtext 1.0pt;padding:0in'>Zygophyseter</span></em>. With its
unique supracranial basin morphology,<span class=apple-converted-space> </span><em><span
style='font-family:"Arial","sans-serif";border:none windowtext 1.0pt;
padding:0in'>Livyatan</span></em><span class=apple-converted-space> </span>forms
a separate, more crownward stem physeteroid lineage. Combined with
biostratigraphic information, our cladistic hypothesis allows us to discuss the
time of origin of the main physeteroid clades, as well as trends in the
evolution of their body size, dentition, temporal fossa and supracranial basin.<o:p></o:p></span></p>

<p class=MsoNormal><o:p> </o:p></p>

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